Gibberellin metabolism and its regulation.

Bioactive gibberellins (GAs) are diterpene plant hormones that are biosynthesized through complex pathways and control diverse aspects of growth and development. Biochemical, genetic, and genomic approaches have led to the identification of the majority of the genes that encode GA biosynthesis and deactivation enzymes. Recent studies have highlighted the occurrence of previously unrecognized deactivation mechanisms. It is now clear that both GA biosynthesis and deactivation pathways are tightly regulated by developmental, hormonal, and environmental signals, consistent with the role of GAs as key growth regulators. In some cases, the molecular mechanisms for fine-tuning the hormone levels are beginning to be uncovered. In this review, I summarize our current understanding of the GA biosynthesis and deactivation pathways in plants and fungi, and discuss how GA concentrations in plant tissues are regulated during development and in response to environmental stimuli.

[1]  W. Schliemann,et al.  Gibberellin conjugates: an overview , 1994, Plant Growth Regulation.

[2]  D. Van Der Straeten,et al.  The plant stress hormone ethylene controls floral transition via DELLA-dependent regulation of floral meristem-identity genes , 2007, Proceedings of the National Academy of Sciences.

[3]  N. Harberd,et al.  Step-by-Step Acquisition of the Gibberellin-DELLA Growth-Regulatory Mechanism during Land-Plant Evolution , 2007, Current Biology.

[4]  Y. Kamiya,et al.  Contribution of gibberellin deactivation by AtGA2ox2 to the suppression of germination of dark-imbibed Arabidopsis thaliana seeds. , 2007, Plant & cell physiology.

[5]  Meimei Xu,et al.  Rice Contains Two Disparate ent-Copalyl Diphosphate Synthases with Distinct Metabolic Functions1 , 2004, Plant Physiology.

[6]  Eunkyoo Oh,et al.  PIL5, a Phytochrome-Interacting bHLH Protein, Regulates Gibberellin Responsiveness by Binding Directly to the GAI and RGA Promoters in Arabidopsis Seeds[W] , 2007, The Plant Cell Online.

[7]  J. B. Reid,et al.  PhyA and cry1 act redundantly to regulate gibberellin levels during de‐etiolation in blue light , 2006 .

[8]  D. Gage,et al.  Molecular Cloning and Photoperiod-Regulated Expression of Gibberellin 20-Oxidase from the Long-Day Plant Spinach , 1996, Plant physiology.

[9]  P. Hedden,et al.  Gibberellin Biosynthesis in Plants and Fungi: A Case of Convergent Evolution? , 2001, Journal of Plant Growth Regulation.

[10]  P. Hedden,et al.  Isolation and Expression of Three Gibberellin 20-Oxidase cDNA Clones from Arabidopsis , 1995, Plant physiology.

[11]  S. Fujioka,et al.  Endogenous Gibberellins from Sporophytes of Two Tree Ferns, Cibotium glaucum and Dicksonia antarctica. , 1988, Plant physiology.

[12]  C. Helliwell,et al.  A plastid envelope location of Arabidopsis ent-kaurene oxidase links the plastid and endoplasmic reticulum steps of the gibberellin biosynthesis pathway. , 2001, The Plant journal : for cell and molecular biology.

[13]  J. Noel,et al.  Methylation of Gibberellins by Arabidopsis GAMT1 and GAMT2[W] , 2007, The Plant Cell Online.

[14]  M. Matsuoka,et al.  Where do gibberellin biosynthesis and gibberellin signaling occur in rice plants? , 2003, The Plant journal : for cell and molecular biology.

[15]  P. Hedden,et al.  Characterization of the Final Two Genes of the Gibberellin Biosynthesis Gene Cluster of Gibberella fujikuroi , 2003, Journal of Biological Chemistry.

[16]  Y. Kamiya,et al.  Light activates the degradation of PIL5 protein to promote seed germination through gibberellin in Arabidopsis. , 2006, The Plant journal : for cell and molecular biology.

[17]  S. Iuchi,et al.  Identification and characterization of Arabidopsis gibberellin receptors. , 2006, The Plant journal : for cell and molecular biology.

[18]  J. B. Reid,et al.  Changes in gibberellin A(1) levels and response during de-etiolation of pea seedlings. , 2000, Plant physiology.

[19]  H. Kawaide,et al.  The gene encoding tobacco gibberellin 3beta-hydroxylase is expressed at the site of GA action during stem elongation and flower organ development. , 1999, The Plant journal : for cell and molecular biology.

[20]  S. Hoffmann-Benning,et al.  On the role of abscisic Acid and gibberellin in the regulation of growth in rice. , 1992, Plant physiology.

[21]  Masatomo Kobayashi,et al.  An Overview of Gibberellin Metabolism Enzyme Genes and Their Related Mutants in Rice1[w] , 2004, Plant Physiology.

[22]  P. Hedden,et al.  The Gibberellin 20-Oxidase of Gibberella fujikuroi Is a Multifunctional Monooxygenase* , 2002, The Journal of Biological Chemistry.

[23]  G. Sandberg,et al.  Arabidopsis KNOXI Proteins Activate Cytokinin Biosynthesis , 2005, Current Biology.

[24]  P. Hedden,et al.  Molecular cloning and functional expression of gibberellin 2- oxidases, multifunctional enzymes involved in gibberellin deactivation. , 1999, Proceedings of the National Academy of Sciences of the United States of America.

[25]  O. Nilsson,et al.  GA4 Is the Active Gibberellin in the Regulation of LEAFY Transcription and Arabidopsis Floral Initiation[W] , 2006, The Plant Cell Online.

[26]  R. Amasino,et al.  Overexpression of a Novel Class of Gibberellin 2-Oxidases Decreases Gibberellin Levels and Creates Dwarf Plants Article, publication date, and citation information can be found at www.plantcell.org/cgi/doi/10.1105/tpc.005975. , 2003, The Plant Cell Online.

[27]  T. Sun,et al.  Synergistic derepression of gibberellin signaling by removing RGA and GAI function in Arabidopsis thaliana. , 2001, Genetics.

[28]  Y. Kamiya,et al.  ELONGATED UPPERMOST INTERNODE Encodes a Cytochrome P450 Monooxygenase That Epoxidizes Gibberellins in a Novel Deactivation Reaction in Rice[W] , 2006, The Plant Cell Online.

[29]  Ayuko Kuwahara,et al.  Gibberellin Biosynthesis and Response during Arabidopsis Seed Germination Online version contains Web-only data. Article, publication date, and citation information can be found at www.plantcell.org/cgi/doi/10.1105/tpc.011650. , 2003, The Plant Cell Online.

[30]  C. Helliwell,et al.  The Involvement of Gibberellin 20-Oxidase Genes in Phytochrome-Regulated Petiole Elongation of Arabidopsis , 2005, Plant Physiology.

[31]  K. Asano,et al.  Molecular Interactions of a Soluble Gibberellin Receptor, GID1, with a Rice DELLA Protein, SLR1, and Gibberellin[W] , 2007, The Plant Cell Online.

[32]  L. T. Evans,et al.  Gibberellins and flowering of grasses and cereals: prizing open the lid of the "florigen" black box. , 2003, Annual review of plant biology.

[33]  J. Ozga,et al.  Seed and Hormonal Regulation of Gibberellin 20-Oxidase Expression in Pea Pericarp , 1997, Plant physiology.

[34]  Patrick Achard,et al.  Integration of Plant Responses to Environmentally Activated Phytohormonal Signals , 2006, Science.

[35]  J. Olsen,et al.  Thermoperiodic Stem Elongation Involves Transcriptional Regulation of Gibberellin Deactivation in Pea1 , 2005, Plant Physiology.

[36]  D. Weigel,et al.  Integration of floral inductive signals in Arabidopsis , 2000, Nature.

[37]  Søren Bak,et al.  Comparative Genomics of Rice and Arabidopsis. Analysis of 727 Cytochrome P450 Genes and Pseudogenes from a Monocot and a Dicot1[w] , 2004, Plant Physiology.

[38]  S. Yamaguchi,et al.  Gibberellin biosynthesis: its regulation by endogenous and environmental signals. , 2000, Plant & cell physiology.

[39]  J. Ozga,et al.  Hormone and Seed-Specific Regulation of Pea Fruit Growth1 , 2002, Plant Physiology.

[40]  S. Yamaguchi,et al.  Distinct cell-specific expression patterns of early and late gibberellin biosynthetic genes during Arabidopsis seed germination. , 2002, The Plant journal : for cell and molecular biology.

[41]  L. T. Evans,et al.  Long-day induction of flowering in Lolium temulentum involves sequential increases in specific gibberellins at the shoot apex. , 2001, Plant physiology.

[42]  M. Koornneef,et al.  Induction and analysis of gibberellin sensitive mutants in Arabidopsis thaliana (L.) heynh. , 1980, Theoretical and Applied Genetics.

[43]  Y. Kamiya,et al.  Contribution of the Mevalonate and Methylerythritol Phosphate Pathways to the Biosynthesis of Gibberellins inArabidopsis * , 2002, The Journal of Biological Chemistry.

[44]  P. Hedden,et al.  Gibberellin biosynthesis and inactivation , 2010 .

[45]  Y. Kamiya,et al.  GIBBERELLIN BIOSYNTHESIS: Enzymes, Genes and Their Regulation. , 1997, Annual review of plant physiology and plant molecular biology.

[46]  J. García-Martínez,et al.  Light regulation of gibberellin A1 content and expression of genes coding for GA 20-oxidase and GA 3β-hydroxylase in etiolated pea seedlings , 2000 .

[47]  J. B. Reid,et al.  Endogenous gibberellin A1 levels control thermoperiodic stem elongation in Pisum sativum , 1998 .

[48]  C. MacMillan,et al.  Flowering of the Grass Lolium perenne. Effects of Vernalization and Long Days on Gibberellin Biosynthesis and Signaling1 , 2005, Plant Physiology.

[49]  H. Kawaide,et al.  Ent-kaurene synthase from the fungus Phaeosphaeria sp. L487. cDNA isolation, characterization, and bacterial expression of a bifunctional diterpene cyclase in fungal gibberellin biosynthesis. , 1997, The Journal of biological chemistry.

[50]  D. Gage,et al.  Feedback Regulation of GA5 Expression and Metabolic Engineering of Gibberellin Levels in Arabidopsis , 1999, Plant Cell.

[51]  J. Ozga,et al.  Specificity of auxin regulation of gibberellin 20-oxidase gene expression in pea pericarp , 2002, Plant Molecular Biology.

[52]  E. Jensen,et al.  Hydrolysis and reconjugation of gibberellin A20 glucosyl ester by seedlings of Zea mays L. , 1992, Proceedings of the National Academy of Sciences of the United States of America.

[53]  Ayuko Kuwahara,et al.  Regulation of hormone metabolism in Arabidopsis seeds: phytochrome regulation of abscisic acid metabolism and abscisic acid regulation of gibberellin metabolism. , 2006, The Plant journal : for cell and molecular biology.

[54]  S. Iwahori,et al.  KNOX homeodomain protein directly suppresses the expression of a gibberellin biosynthetic gene in the tobacco shoot apical meristem. , 2001, Genes & development.

[55]  C. Helliwell,et al.  Cloning of the Arabidopsis ent-kaurene oxidase gene GA3. , 1998, Proceedings of the National Academy of Sciences of the United States of America.

[56]  Xiaoying Zhao,et al.  A Study of Gibberellin Homeostasis and Cryptochrome-Mediated Blue Light Inhibition of Hypocotyl Elongation1[W][OA] , 2007, Plant Physiology.

[57]  Y. Kamiya,et al.  Emission of ent-kaurene, a diterpenoid hydrocarbon precursor for gibberellins, into the headspace from plants. , 2004, Plant & cell physiology.

[58]  M. Matsuoka,et al.  Over-expression of a tobacco homeobox gene, NTH15, decreases the expression of a gibberellin biosynthetic gene encoding GA 20-oxidase. , 1998, The Plant journal : for cell and molecular biology.

[59]  S. Tabata,et al.  Distinct and overlapping roles of two gibberellin 3-oxidases in Arabidopsis development. , 2006, The Plant journal : for cell and molecular biology.

[60]  Jennifer J. Smith,et al.  Auxin from the Developing Inflorescence Is Required for the Biosynthesis of Active Gibberellins in Barley Stems1 , 2004, Plant Physiology.

[61]  M. Matsuoka,et al.  Cloning and functional analysis of two gibberellin 3β-hydroxylase genes that are differently expressed during the growth of rice , 2001, Proceedings of the National Academy of Sciences of the United States of America.

[62]  L. T. Evans,et al.  Regulation of Flowering in the Long-Day Grass Lolium temulentum by Gibberellins and the FLOWERING LOCUS T Gene , 2006, Plant Physiology.

[63]  P. Hedden,et al.  Gibberellin metabolism: new insights revealed by the genes. , 2000, Trends in plant science.

[64]  P. Saxena,et al.  A Crucial Role for Gibberellins in Stress Protection of Plants , 1999 .

[65]  S. Ishida,et al.  AGF1, an AT-Hook Protein, Is Necessary for the Negative Feedback of AtGA3ox1 Encoding GA 3-Oxidase1[W] , 2007, Plant Physiology.

[66]  T. Toyomasu,et al.  Biological functions of ent- and syn-copalyl diphosphate synthases in rice: key enzymes for the branch point of gibberellin and phytoalexin biosynthesis. , 2004, The Plant journal : for cell and molecular biology.

[67]  F. Parcy,et al.  AtGA3ox2, a Key Gene Responsible for Bioactive Gibberellin Biosynthesis, Is Regulated during Embryogenesis by LEAFY COTYLEDON2 and FUSCA3 in Arabidopsis1 , 2004, Plant Physiology.

[68]  J. Ross,et al.  Auxin Regulation of the Gibberellin Pathway in Pea1 , 2002, Plant Physiology.

[69]  D. Alabadí,et al.  Transcriptional Regulation of Gibberellin Metabolism Genes by Auxin Signaling in Arabidopsis1[W] , 2006, Plant Physiology.

[70]  Tai-Ping Sun,et al.  Gibberellin signaling: biosynthesis, catabolism, and response pathways. , 2002, The Plant cell.

[71]  D. J. Lee,et al.  Differential Regulation of RNA Levels of Gibberellin Dioxygenases by Photoperiod in Spinach1 , 2002, Plant Physiology.

[72]  D. J. Lee,et al.  Molecular Cloning of GA 2-Oxidase3 from Spinach and Its Ectopic Expression in Nicotiana sylvestris1 , 2005, Plant Physiology.

[73]  I. Yamaguchi,et al.  Identification of Ten Gibberellins from Sporophytes of the Tree fern, Cyathea australis. , 1985, Plant physiology.

[74]  M. Nishimura,et al.  Reduction to below threshold levels of glycolate oxidase activities in transgenic tobacco enhances photoinhibition during irradiation. , 2000, Plant & cell physiology.

[75]  K. Halliday,et al.  Cold and Light Control Seed Germination through the bHLH Transcription Factor SPATULA , 2005, Current Biology.

[76]  Masatomo Kobayashi,et al.  The dwarf-1 (dt) Mutant of Zea mays blocks three steps in the gibberellin-biosynthetic pathway. , 1996, Proceedings of the National Academy of Sciences of the United States of America.

[77]  Joan Gil,et al.  Light Regulation of Gibberellin Biosynthesis and Mode of Action , 2001, Journal of Plant Growth Regulation.

[78]  M. Matsuoka,et al.  Alteration of hormone levels in transgenic tobacco plants overexpressing the rice homeobox gene OSH1. , 1998, Plant physiology.

[79]  R. Peters,et al.  Synergistic Substrate Inhibition of ent-Copalyl Diphosphate Synthase: A Potential Feed-Forward Inhibition Mechanism Limiting Gibberellin Metabolism1[OA] , 2007, Plant Physiology.

[80]  S. Hake,et al.  The Gibberellin Pathway Mediates KNOTTED1-Type Homeobox Function in Plants with Different Body Plans , 2002, Current Biology.

[81]  P. Hedden,et al.  Modification of gibberellin production and plant development in Arabidopsis by sense and antisense expression of gibberellin 20-oxidase genes. , 1999, The Plant journal : for cell and molecular biology.

[82]  Yuji Kamiya,et al.  PHYTOCHROME REGULATION OF GIBBERELLIN 3β-HYDROXYLASE GENES IN GERMINATING ARABIDOPSIS SEEDS , 1999 .

[83]  T. Shinomura,et al.  Action spectra for phytochrome A- and B-specific photoinduction of seed germination in Arabidopsis thaliana. , 1996, Proceedings of the National Academy of Sciences of the United States of America.

[84]  T. Sun,et al.  Molecular mechanism of gibberellin signaling in plants. , 2004, Annual review of plant biology.

[85]  Masatomo Kobayashi,et al.  Expression of novel rice gibberellin 2-oxidase gene is under homeostatic regulation by biologically active gibberellins , 2003, Journal of plant research.

[86]  Y. Kamiya,et al.  REPRESSION OF SHOOT GROWTH, a bZIP Transcriptional Activator, Regulates Cell Elongation by Controlling the Level of Gibberellins , 2000, Plant Cell.

[87]  P. Hedden,et al.  Function and transcript analysis of gibberellin-biosynthetic enzymes in wheat , 2006, Planta.

[88]  S. Yamaguchi,et al.  The GA2 locus of Arabidopsis thaliana encodes ent-kaurene synthase of gibberellin biosynthesis. , 1998, Plant physiology.

[89]  D. Alabadí,et al.  Gibberellins Repress Photomorphogenesis in Darkness1 , 2004, Plant Physiology.

[90]  H. Kawaide,et al.  Cloning of a Full-length cDNA Encoding ent-Kaurene Synthase from Gibberella fujikuroi: Functional Analysis of a Bifunctional Diterpene Cyclase , 2000, Bioscience, biotechnology, and biochemistry.

[91]  T. Shinomura Phytochrome regulation of seed germination , 1997, Journal of Plant Research.

[92]  M. Matsuoka,et al.  A Rice Semi-Dwarf Gene, Tan-Ginbozu (D35), Encodes the Gibberellin Biosynthesis Enzyme, ent-Kaurene Oxidase , 2004, Plant Molecular Biology.

[93]  P. Hedden,et al.  KNOX Action in Arabidopsis Is Mediated by Coordinate Regulation of Cytokinin and Gibberellin Activities , 2005, Current Biology.

[94]  H. Kawaide,et al.  Cloning and characterization of a cDNA encoding gibberellin 20‐oxidase from rice (Oryza sativa) seedlings , 1997 .

[95]  Masatomo Kobayashi,et al.  Accumulation of Phosphorylated Repressor for Gibberellin Signaling in an F-box Mutant , 2003, Science.

[96]  Meimei Xu,et al.  Functional identification of rice syn-copalyl diphosphate synthase and its role in initiating biosynthesis of diterpenoid phytoalexin/allelopathic natural products. , 2004, The Plant journal : for cell and molecular biology.

[97]  D. Robinson,et al.  ent-Kaurene synthase is located in proplastids of meristematic shoot tissues , 1997, Planta.

[98]  T. Sun,et al.  Overexpression of AtCPS and AtKS in Arabidopsis Confers Increased ent-Kaurene Production But No Increase in Bioactive Gibberellins1 , 2003, Plant Physiology.

[99]  I. Hwang,et al.  Isolation of the Arabidopsis GA4 locus. , 1995, The Plant cell.

[100]  S. Iwahori,et al.  Expression of a gibberellin 2-oxidase gene around the shoot apex is related to phase transition in rice. , 2001, Plant physiology.

[101]  L. T. Evans,et al.  Synthesis of gibberellin GA6 and its role in flowering of Lolium temulentum. , 2003, Phytochemistry.

[102]  Y. Kamiya,et al.  Functional Analysis of the Two Interacting Cyclase Domains in ent-Kaurene Synthase from the FungusPhaeosphaeria sp. L487 and a Comparison with Cyclases from Higher Plants* , 2000, The Journal of Biological Chemistry.

[103]  R. Foster,et al.  Control of specific gene expression by gibberellin and brassinosteroid. , 2001, Plant physiology.

[104]  T. Sun,et al.  The Arabidopsis GA1 locus encodes the cyclase ent-kaurene synthetase A of gibberellin biosynthesis. , 1994, The Plant cell.

[105]  K. Oda,et al.  dwarf and delayed-flowering 1, a novel Arabidopsis mutant deficient in gibberellin biosynthesis because of overexpression of a putative AP2 transcription factor. , 2004, The Plant journal : for cell and molecular biology.

[106]  J. B. Reid,et al.  The brassinosteroid growth response in pea is not mediated by changes in gibberellin content , 2005, Planta.

[107]  M. Matsuoka,et al.  The α-Amylase Induction in Endosperm during Rice Seed Germination Is Caused by Gibberellin Synthesized in Epithelium1 , 2002, Plant Physiology.

[108]  J. B. Reid,et al.  Abscisic acid levels in seeds of the gibberellin-deficient mutant lh-2 of pea (Pisum sativum) , 1999 .

[109]  Yuji Kamiya,et al.  Activation of Gibberellin Biosynthesis and Response Pathways by Low Temperature during Imbibition of Arabidopsis thaliana Seeds On-line version contains Web-only data. , 2004, The Plant Cell Online.

[110]  H. Kawaide Biochemical and Molecular Analyses of Gibberellin Biosynthesis in Fungi , 2006, Bioscience, biotechnology, and biochemistry.

[111]  Yu Zhao,et al.  The Rice YABBY1 Gene Is Involved in the Feedback Regulation of Gibberellin Metabolism1[C][W] , 2007, Plant Physiology.

[112]  D. J. Lee,et al.  Regulation of gibberellin 20-oxidase1 expression in spinach by photoperiod , 2007, Planta.

[113]  H. Kawaide,et al.  Gibberellin biosynthesis in Gibberella fujikuroi: cloning and characterization of the copalyl diphosphate synthase gene , 1998, Current Genetics.

[114]  Huai Wang,et al.  The Embryo MADS Domain Protein AGAMOUS-Like 15 Directly Regulates Expression of a Gene Encoding an Enzyme Involved in Gibberellin Metabolism , 2004, The Plant Cell Online.

[115]  J. B. Reid,et al.  Regulation of gibberellin 20-oxidase and gibberellin 3beta-hydroxylase transcript accumulation during De-etiolation of pea seedlings. , 1999, Plant physiology.

[116]  J. Macmillan Occurrence of Gibberellins in Vascular Plants, Fungi, and Bacteria , 2001, Journal of Plant Growth Regulation.

[117]  Masatomo Kobayashi,et al.  GIBBERELLIN INSENSITIVE DWARF1 encodes a soluble receptor for gibberellin , 2005, Nature.

[118]  H. Kawaide,et al.  Identification and functional analysis of bifunctional ent‐kaurene synthase from the moss Physcomitrella patens , 2006, FEBS letters.

[119]  R. Quatrano,et al.  Mosses as model systems for the study of metabolism and development. , 2006, Annual review of plant biology.

[120]  H. Kawaide,et al.  Deactivation of Gibberellin by 2-Oxidation during Germination of Photoblastic Lettuce Seeds , 2003, Bioscience, biotechnology, and biochemistry.

[121]  Carla M. Wolbang,et al.  Auxin promotes gibberellin biosynthesis in decapitated tobacco plants , 2001, Planta.

[122]  H. Kawaide,et al.  Phytochrome regulates gibberellin biosynthesis during germination of photoblastic lettuce seeds. , 1998, Plant physiology.

[123]  M. Matsuoka,et al.  Ectopic Expression of KNOTTED1-Like Homeobox Protein Induces Expression of Cytokinin Biosynthesis Genes in Rice1[W] , 2006, Plant Physiology.

[124]  Gerco C. Angenent,et al.  Transcriptional program controlled by the floral homeotic gene AGAMOUS during early organogenesis , 2005, Development.

[125]  C. Helliwell,et al.  The CYP88A cytochrome P450, ent-kaurenoic acid oxidase, catalyzes three steps of the gibberellin biosynthesis pathway. , 2001, Proceedings of the National Academy of Sciences of the United States of America.

[126]  P. Hedden,et al.  The P450–1 gene of Gibberella fujikuroi encodes a multifunctional enzyme in gibberellin biosynthesis , 2001, Proceedings of the National Academy of Sciences of the United States of America.

[127]  P. Hedden,et al.  Gibberellins and seed development in maize. I. Evidence that gibberellin/abscisic acid balance governs germination versus maturation pathways. , 2000, Plant physiology.

[128]  Masanori Okamoto,et al.  The transcription factor FUSCA3 controls developmental timing in Arabidopsis through the hormones gibberellin and abscisic acid. , 2004, Developmental cell.

[129]  J. B. Reid,et al.  Hormone Distribution and Transport , 2007 .

[130]  J. Smith,et al.  Evidence that auxin promotes gibberellin A1 biosynthesis in pea. , 2000, The Plant journal : for cell and molecular biology.

[131]  Shihshieh Huang,et al.  Overexpression of 20-oxidase confers a gibberellin-overproduction phenotype in Arabidopsis. , 1998, Plant Physiology.

[132]  M. Talón,et al.  Gibberellin A1 is required for stem elongation in spinach. , 1993, Proceedings of the National Academy of Sciences of the United States of America.

[133]  T. Moritz,et al.  Gibberellins are not required for normal stem growth in Arabidopsis thaliana in the absence of GAI and RGA. , 2001, Genetics.

[134]  S. Ishida,et al.  14-3-3 Proteins Regulate Intracellular Localization of the bZIP Transcriptional Activator RSG Article, publication date, and citation information can be found at www.aspb.org/cgi/doi/10.1105/tpc.010188. , 2001, The Plant Cell Online.

[135]  T. Sun,et al.  Developmental regulation of the gibberellin biosynthetic gene GA1 in Arabidopsis thaliana. , 1997, The Plant journal : for cell and molecular biology.

[136]  M. Matsuoka,et al.  Gibberellin receptor and its role in gibberellin signaling in plants. , 2007, Annual review of plant biology.

[137]  T. Sun,et al.  Regulation and cellular localization of ent‐kaurene synthesis , 1997 .

[138]  S. Ishida,et al.  Involvement of 14-3-3 Signaling Protein Binding in the Functional Regulation of the Transcriptional Activator REPRESSION OF SHOOT GROWTH by Gibberellins , 2004, The Plant Cell Online.

[139]  D. Weiss,et al.  Identification of Endogenous Gibberellins in Petunia Flowers (Induction of Anthocyanin Biosynthetic Gene Expression and the Antagonistic Effect of Abscisic Acid) , 1995, Plant physiology.

[140]  M. Ellis,et al.  Semidwarf (sd-1), “green revolution” rice, contains a defective gibberellin 20-oxidase gene , 2002, Proceedings of the National Academy of Sciences of the United States of America.

[141]  J. B. Reid,et al.  Control of Gibberellin Levels and Gene Expression during De-Etiolation in Pea1 , 2002, Plant Physiology.

[142]  D. Gage,et al.  Gibberellins and Stem Growth in Arabidopsis thaliana (Effects of Photoperiod on Expression of the GA4 and GA5 Loci) , 1997, Plant physiology.