Role Of Gibberellins Research Articles

Long-distance regulation of flowering time

One of the great mysteries of plant science appears to have been resolved with the discovery that the protein FT can act as a phloem-mobile florigen hormone. The collective evidence from several laboratories, many from studies on photoperiod response, indicates that FT and its homologues are universal signalling molecules for flowering plants. Duplication and divergence of FT-like proteins reveals an increased complexity of function in certain taxonomic groups including grasses and legumes. There are additional components of long-distance flowering time control, such as a role for gibberellins in some species but probably not others. Cytokinins and sugars are further putative signals. Vernalization processes and responses are generally considered to occur in shoot meristems, but systemic responses to cold have been reported several times. Finally, there is increasing evidence that FT does not act purely to switch on flowering, but in addition, has broader roles in seasonal developmental switches such as bud dormancy and tuberization, and in the regulation of meristem determinacy and compound leaf development. This review seeks to highlight recent progress in systemic floral signalling, and to indicate areas in need of further research.

Occurrence of plant growth regulators in Psilotum nudum

The endogenous content of the auxin indole-3-acetic acid (IAA), the cytokinins trans-zeatin (tZ), trans-zeatin riboside (tZR), dihydrozeatin (DHZ), dihydrozeatin riboside (DHZR), isopentenyladenine (iP) and isopentenyladenosine (iPR), the gibberellins GA 1, GA 3, GA 4, GA 7, GA 9 and GA 20 in the rhizome and aerial shoots during and after sporogenesis were measured by high-performance liquid chromatography–tandem mass spectrometry in the fern Psilotum nudum. The present study shows, for the first time, the presence of the auxin IAA, the cytokinins tZR, DHZR and iP, and the gibberellins GA 4, GA 9 and GA 20 in the rhizome and aerial shoots of this species and suggests a possible role of gibberellins in the evolution of ferns.

Proteome study of the phloem sap of pumpkin using multidimensional protein identification technology

The phloem is the major transport route for both small substances and large molecules, such as proteins and RNAs, from their sources to sink tissues. To investigate the proteins present in pumpkin phloem sap, proteome analysis using multidimensional protein identification technology was carried out. Pumpkin phloem peptides obtained by liquid chromatography/mass spectrometry/mass spectrometry were searched against pumpkin protein data derived from the National Center for Biotechnology Information. A total of 47 pumpkin phloem proteins were identified. The identified proteins mainly corresponded to enzymes involved in gibberellin biosynthesis, antioxidation processes, or defense mechanisms. Interestingly, seven enzymes required for gibberellin biosynthesis were identified for the first time by this proteomics approach. In summary, the new phloem proteins identified in this study provide strong evidence for stress and defense signaling and new insights regarding the role of gibberellin in the developmental programming of higher plants through the phloem.

A survey of flowering genes reveals the role of gibberellins in floral control in rose

Exhaustive studies on flowering control in annual plants have provided a framework for exploring this process in other plant species, especially in perennials for which little molecular data are currently available. Rose is a woody perennial plant with a particular flowering strategy--recurrent blooming, which is controlled by a recessive locus (RB). Gibberellins (GA) inhibit flowering only in non-recurrent roses. Moreover, the GA content varies during the flowering process and between recurrent and non-recurrent rose. Only a few rose genes potentially involved in flowering have been described, i.e. homologues of ABC model genes and floral genes from EST screening. In this study, we gained new information on the molecular basis of rose flowering: date of flowering and recurrent blooming. Based on a candidate gene strategy, we isolated genes that have similarities with genes known to be involved in floral control in Arabidopsis (GA pathway, floral repressors and integrators). Candidate genes were mapped on a segregating population, gene expression was studied in different organs and transcript abundance was monitored in growing shoot apices. Twenty-five genes were studied. RoFT, RoAP1 and RoLFY are proposed to be good floral markers. RoSPY and RB co-localized in our segregating population. GA metabolism genes were found to be regulated during floral transition. Furthermore, GA signalling genes were differentially regulated between a non-recurrent rose and its recurrent mutant. We propose that flowering gene networks are conserved between Arabidopsis and rose. The GA pathway appears to be a key regulator of flowering in rose. We postulate that GA metabolism is involved in floral initiation and GA signalling might be responsible for the recurrent flowering character.

Gibberellin controls the nodulation signaling pathway in Lotus japonicus

Root nodule formation is regulated by several plant hormones, but the details of the regulation of the nodulation signaling pathway are largely unknown. In this study, the role of gibberellin (GA) in the control of root nodule symbiosis was investigated at the physiological and genetic levels in Lotus japonicus. Exogenous application of biologically active GA, GA(3), inhibited the formation of infection threads and nodules, which was counteracted by the application of a biosynthesis inhibitor of GA, Uniconazole P. Nod factor-induced root hair deformation was severely blocked in the presence of GA, which was phenocopied by nsp2 mutants. The number of spontaneous nodules triggered by the gain-of-function mutation of calcium/calmodulin-dependent kinase (CCaMK) or the lotus histidine kinase 1 (LHK1) was decreased upon the addition of GA; moreover, the overexpression of the gain-of-function mutation of L. japonicus, SLEEPY1, a positive regulator of GA signaling, resulted in a reduced nodule number, without other aspects of root development being affected. These results indicate that higher GA signaling levels specifically inhibit the nodulation signaling pathway. Nod factor-dependent induction of NSP2 and NIN was inhibited by exogenous GA. Furthermore, the cytokinin-dependent induction of NIN was suppressed by GA. From these results, we conclude that GA inhibits the nodulation signaling pathway downstream of cytokinin, possibly at NSP2, which is required for Nod factor-dependent NIN expression. These results clarify the roles of GA in the nodulation signaling pathway, and in relation to the cytokinin signaling pathway for nodulation in L. japonicus.

Gibberellins regulate lateral root formation in Populus through interactions with auxin and other hormones.

The role of gibberellins (GAs) in regulation of lateral root development is poorly understood. We show that GA-deficient (35S:PcGA2ox1) and GA-insensitive (35S:rgl1) transgenic Populus exhibited increased lateral root proliferation and elongation under in vitro and greenhouse conditions, and these effects were reversed by exogenous GA treatment. In addition, RNA interference suppression of two poplar GA 2-oxidases predominantly expressed in roots also decreased lateral root formation. GAs negatively affected lateral root formation by inhibiting lateral root primordium initiation. A whole-genome microarray analysis of root development in GA-modified transgenic plants revealed 2069 genes with significantly altered expression. The expression of 1178 genes, including genes that promote cell proliferation, growth, and cell wall loosening, corresponded to the phenotypic severity of the root traits when transgenic events with differential phenotypic expression were compared. The array data and direct hormone measurements suggested crosstalk of GA signaling with other hormone pathways, including auxin and abscisic acid. Transgenic modification of a differentially expressed gene encoding an auxin efflux carrier suggests that GA modulation of lateral root development is at least partly imparted by polar auxin transport modification. These results suggest a mechanism for GA-regulated modulation of lateral root proliferation associated with regulation of plant allometry during the stress response.

Temperature perception and signal transduction in plants

Plants can show remarkable responses to small changes in temperature, yet one of the great unknowns in plant science is how that temperature signal is perceived. The identity of the early components of the temperature signal transduction pathway also remains a mystery. To understand the consequences of anthropogenic environmental change we will have to learn much more about the basic biology of how plants sense temperature. Recent advances show that many known plant-temperature responses share common signalling components, and suggest ways in which these might be linked to form a plant temperature signalling network.

Roles of gibberellins and abscisic acid in dormancy and germination of red bayberry (Myrica rubra) seeds

Intact seeds from freshly harvested fruits of Myrica rubra (Sieb et Zucc.) were dormant and required 8 weeks of warm stratification followed by 12 weeks of cold stratification for germination. Exogenous application of gibberellic acid (GA(3)) to intact fresh seeds was effective in breaking dormancy, with > 70% of seeds germinating when treated with 5.2 mM GA(3) and incubated at a day/night temperature of 30/20 degrees C for 20 weeks. Removing the hard endocarp or endocarp plus seed coat of fresh seeds promoted germination, and addition of GA(3) to the embryo accelerated germination. The gibberellins GA(1) and GA(4) were more effective than GA(3) in promoting germination of seeds with the endocarp removed. Endogenous contents of GA(1), GA(3), GA(4), GA(7) and GA(20) were quantified by gas chromatography-mass spectrometry-selected ion monitoring in the endocarps, seed coats and embryos of fresh seeds treated with 5.2 mM GA(3). The content of GA(3) decreased in the endocarp during incubation, whereas GA(1) contents increased in the endocarp and seed coat. A high GA(1) content was detected in the endocarps and embryos of newly germinated seeds. We speculate that GA(3) was converted to GA(1) during incubation and that GA(1) is involved in seed germination. Endogenous abscisic acid (ABA) contents were measured in fresh seeds and in warm and cold stratified seeds. The ABA content in fresh seeds was distributed in the order endocarp > seed coat > embryo, with the content in the endocarp being about 132-fold higher than in the seed coat and embryo. Total ABA content of seeds subjected to warm or cold stratification, or both, was 8.7- to 14.0-fold lower than that of fresh seeds. Low contents of endogenous GA(1), GA(3), GA(7) and GA(20), but elevated contents of GA(4), were found in the seed coats and endocarps of warm plus cold stratified seeds and in the seed coats and embryos of newly germinated seeds. These observations, coupled with the finding that GA stimulated germination of dormant Myrica seeds, provide evidence that endogenous ABA inhibited release of dormancy and that endogenous gibberellins, especially GA(4) or GA(1), or both, are involved in germination.

Effect of gibberellin on Arabidopsis growth, development, and hormonal balance under green and blue light

The role of gibberellins in plant morphology under selective light was studied. A comparison of the effects of green and blue light on growth, development, and hormonal balance was performed for two Arabidopsis thaliana (L.) Heynh. (ecotype Landsberg erecta lines: wild type Ler and its ga4-1 mutant with suppressed GA4/1 synthesis. The absence of active GA4/1 from ga4-1 mutant determined its retarded growth, slowed passing through developmental phases, suppressed apical dominance, and reduced seed productivity. The retarded growth and development of the mutant was related to changed hormonal balance in them as compared to wild-type line: IAA content and the IAA/ABA ratio were declined, zeatin riboside and ABA accumulated. Green light retarded stem elongation and branching, reduced leaf specific surface density and plant seed productivity, and retarded plant transition to reproduction to a greater degree at GA4/1 deficit (ga4-1) than at its normal content (Ler).

Gibberellin homeostasis and plant height control by EUI and a role for gibberellin in root gravity responses in rice

The rice Eui (ELONGATED UPPERMOST INTERNODE) gene encodes a cytochrome P450 monooxygenase that deactivates bioactive gibberellins (GAs). In this study, we investigated controlled expression of the Eui gene and its role in plant development. We found that Eui was differentially induced by exogenous GAs and that the Eui promoter had the highest activity in the vascular bundles. The eui mutant was defective in starch granule development in root caps and Eui overexpression enhanced starch granule generation and gravity responses, revealing a role for GA in root starch granule development and gravity responses. Experiments using embryoless half-seeds revealed that RAmy1A and GAmyb were highly upregulated in eui aleurone cells in the absence of exogenous GA. In addition, the GA biosynthesis genes GA3ox1 and GA20ox2 were downregulated and GA2ox1 was upregulated in eui seedlings. These results indicate that EUI is involved in GA homeostasis, not only in the internodes at the heading stage, but also in the seedling stage, roots and seeds. Disturbing GA homeostasis affected the expression of the GA signaling genes GID1 (GIBBERELLIN INSENSITIVE DWARF 1), GID2 and SLR1. Transgenic RNA interference of the Eui gene effectively increased plant height and improved heading performance. By contrast, the ectopic expression of Eui under the promoters of the rice GA biosynthesis genes GA3ox2 and GA20ox2 significantly reduced plant height. These results demonstrate that a slight increase in Eui expression could dramatically change rice morphology, indicating the practical application of the Eui gene in rice molecular breeding for a high yield potential.

End-of-day far-red treatment enhances responsiveness to gibberellins and promotes stem extension in chrysanthemum

SUMMARYA brief end-of-day (EOD) far-red (FR) exposure promotes extension growth in plants. This change suggests a role for gibberellins (GA). For chrysanthemum (Chrysanthemum morifolium Ramat.), we show that EOD-FR exposure enhances shoot extension and is mediated, at least in part, by increased responsiveness (sensitivity) to GA. In addition, our analysis showed how light, especially EOD-FR, influenced shoot extension. The effect of EOD-FR on promoting extension growth was maintained, not only during the dark period, but also in the subsequent light period. Furthermore, our observations suggest that dark reversion of certain phytochromes is involved in the determination of flowering time. We also discuss the applicability of EOD-FR treatment to reduce the overall duration of cultivation, while maintaining a sufficient stem height during cut chrysanthemum production.

Roles of gibberellins in increasing sink demand in Japanese pear fruit during rapid fruit growth

Our previous work demonstrated that exogenous gibberellins (GAs) applications during rapid fruit growth significantly increases sink demand and results in a larger fruit in Japanese pear. In an attempt to unravel the mechanism of increased sink demand by applied GAs, the histology, cell wall components of the flesh, and carbon accumulation in the fruit were assessed for Japanese pear (Pyrus pyrifolia, cultivar ‘Kousui’), as were the activities of sucrose- and sorbitol-cleaving enzymes. Our results show that most vascular tissues occurred in core tissue with very little vascular tissue in the flesh. Application of a mixture of GA3 + GA4 in lanolin paste significantly increased the amount of ethanol-insoluble solids, e.g., total pectins, hemicellulose, and cellulose in the cell walls. There was a significantly increased sink demand (assessed by 13C accumulation in the fruit) by the applied GAs, and this increased sink strength was closely related to increased activities of cell wall-bound invertase in the core, neutral invertase and NAD-dependent sorbitol dehydrogenase in the flesh during rapid fruit growth. As well, concentrations of sorbitol and sucrose in the flesh were decreased by GA application, while glucose concentration increased. Most importantly, the fact that sink activity can be increased by GA application implies that endogenous GAs are likely to be important modulators for sugar metabolism. Hence, selecting for genotypes with elevated GA production in the growing fruit and increased activities of key enzymes for sugar metabolism could result in increased fruit size.

Flowering in Tobacco Needs Gibberellins but is not Promoted by the Levels of Active GA1 and GA4 in the Apical Shoot

Flowering of Nicotiana tabacum cv Xhanti depends on gibberellins because gibberellin-deficient plants, due to overexpression of a gibberellin 2-oxidase gene (35S:NoGA2ox3) or to treatment with the gibberellin biosynthesis inhibitor paclobutrazol, flowered later than wild type. These plants also showed inhibition of the expression of molecular markers related to floral transition (NtMADS-4 and NtMADS-11). To investigate further the role of gibberellin in flowering, we quantified its content in tobacco plants during development. We found a progressive reduction in the levels of GA1 and GA4 in the apical shoot during vegetative growth, reaching very low levels at floral transition and beyond. This excludes these two gibberellins as flowering-promoting factors in the apex. The evolution of active gibberellin content in apical shoots agrees with the expression patterns of gibberellin metabolism genes: two encoding gibberellin 20-oxidases (NtGA20ox1 = Ntc12, NtGA20ox2 = Ntc16), one encoding a gibberellin 3-oxidase (NtGA3ox1 = Nty) and one encoding a gibberellin 2-oxidase (NtGA2ox1), suggesting that active gibberellins are locally synthesized. In young apical leaves, GA1 and GA4 content and the expression of gibberellin metabolism genes were rather constant. Our results support that floral transition in tobacco, in contrast to that in Arabidopsis, is not regulated by the levels of GA1 and GA4 in apical shoots, although reaching a threshold in gibberellin levels may be necessary to allow meristem competence for flowering.

Gibberellins and antheridiogen on sex in Blechnum spicant L.

The role of gibberellins (GAs) in determining sex in the gametophyte of the fern Blechnum spicant L. was studied through (a) the effect of exogenous GA(4+7) and GA3 (b) quantitation of the endogenous levels of GA1, GA3, GA4, GA7, GA9, and GA20 in male and female gametophytes, and (c) the effect of flurprimidol, a GAs biosynthesis inhibitor of the steps of oxidation of ent-kaureno to ent-kaurenoic acid. Our results show that GA(4+7) had a slight effect of inducing either male or female sexual organs, antheridia and archegonia, respectively. The endogenous GAs content was not significantly different between sexes, but the GA4, GA7, and GA20 levels were raised above those of the other GAs in both sexes. Neither antheridiogen biosynthesis nor antheridia formation was inhibited by flurprimidol. Gametophytes regenerated from homogenized mature gametophytes (HG) show a different physiological behavior than spore-derived gametophytes. In the first case, gametophytes are males and synthesize antheridiogen before they attain maturity, in contrast to what occurs in spore-derived gametophytes which are females and synthesize antheridiogen when mature.

Exogenous and endogenous growth regulators on apogamy in Dryopteris affinis (Lowe) Fraser-Jenkins sp. affinis

This work showed for the first time the relationship between the effect of exogenous auxins and gibberellins on apogamy in Dryopteris affinis (Lowe) Fraser-Jenkins sp. affinis and its endogenous contents during early apogamic events. The addition of NAA (0.53 and 5.37 microM) or GA(3) 2.8 microM to an MS solid medium significantly increased apogamous sporophyte formation. BA induced brown callus that regenerated sporophytes in a hormone-free medium. The endogenous contents of GA(1), GA(3), GA(4), GA(7), GA(9) and IAA were determined by GC-MS in gametophytes cultured on MS solid medium, before and during early stages of apogamous embryo development. The accumulation of both GA(9) and IAA before embryo development was evident as high levels of GA(4) in the earliest analysed stage of embryo development and high levels of GA(3) in elongating shoots were found. The role of gibberellins on apogamy was also supported by data showing a decrease in the percentage of gametophytes developing embryos because of the addition of flurprimidol to the culture medium.

Inhibitory role of gibberellins in theobroxide-induced flowering of Pharbitis nil

Theobroxide, a novel active compound isolated from a fungus, has been reported previously to induce potato tuberization and flower bud formation in Pharbitis nil under non-inductive long-day conditions. Up to date, the action mechanism of theobroxide on flower-bud induction of P. nil, however, is still unknown. In the present study, we observed a reduction of the stem length, along with the induction of flower buds, in theobroxide-treated and short-day-grown P. nil plants. Also, the results showed that flower bud formation was delayed markedly in P. nil seedlings with removal of cotyledons or exposure to night break. The suppression effect of night-break and cotyledon-removal, however, was abolished completely by spraying theobroxide. Endogenous gibberellin(1/3) contents in P. nil plants treated with theobroxide or grown under short-day conditions were relatively lower, suggesting that gibberellins probably are negatively involved in theobroxide- and short-day-induced flower-bud formation of P. nil.

A DELLAcate balance: the role of gibberellin in plant morphogenesis

The importance of gibberellin (GA) in vegetative and reproductive development has been known for some time. Recent studies have uncovered new roles of GA in leaf differentiation, photomorphogenesis and pollen-tube growth. Significant contributions to our understanding of GA-regulated morphogenesis include the identification of upstream regulators of GA biosynthesis, the elucidation of the function of GA signaling components, and the isolation of downstream targets. In addition, the mechanisms of interactions between GA and other hormone pathways are beginning to be revealed at the molecular level.

Effects of applied gibberellic acid and paclobutrazol on leaf expansion and biomass allocation in two Aegilops species with contrasting leaf elongation rates

The role of gibberellin (GA) in leaf elongation has long been known, however, its involvement in whole shoot growth and biomass allocation is much less clear. We studied the effects of exogenously supplied GA3 and paclobutrazol, an inhibitor of GA biosynthesis, on these processes in Aegilops caudata and Aegilops tauschii, species with contrasting leaf growth characteristics. In both species, addition of GA3 increased leaf elongation rate (LER) through its promoting effect on both cell size and cell number, while paclobutrazol decreased it. Similarly, GA3 increased biomass allocation to the leaves, mainly leaf sheaths, at the cost of allocation to the roots, whereas paclobutrazol had the opposite effect in both species. Despite the increase in LER and biomass allocation to the shoot upon GA3 application, the relative growth rate (RGR) remained constant. Specific leaf area (SLA) was only temporarily affected by GA3 addition. Our results show that the inherent differences in LER and biomass allocation between the slow‐elongating A. caudata and the fast‐elongating A. tauschii are considerably reduced by the exogenous supply of GA3 to the slow‐elongating species, or paclobutrazol to the fast‐elongating one. This suggests a role for gibberellins in explaining inherent differences in leaf area expansion and biomass allocation between the two species in this study.

Effect of Light Source and Quality on Endogenous Gibberellin Level and GA3 Response of Petunia (Petunia*hybrida Vilm.)

Petunia (Petunia X hybrida Vilm.) 'Baccarat Blue' under high pressure sodium lamp (HPS) exhibits, dwarfism compared with one grown under a daylight type metal halide lamp (MH). The role of gibberellin (GA) in this growth change, caused by artificial lighting was investigated. The amounts of GA20 and GA1 were lower in petunia grown under HPS than those under MH, but, the levels of GA19 were similar in both treatments. Plant height of petunia was decreased by treatment with uniconazol, but the difference in plant height between treatments was small. GA3 application promoted stem elongation of uniconazol-treated petunia; the magnitude of stem elongation was independent of light source. These results indicate that dwarfism in petunia under HPS is caused by lower GA20 and GA1 contents, but not by GA sensitivity. The GA biosynthesis step of GA19 to GA20 may be affected by the light quality of these lamps.

The HECT ubiquitin-protein ligase (UPL) family in Arabidopsis: UPL3 has a specific role in trichome development.

Attachment of one or more ubiquitins (Ubs) to various intracellular proteins has a number of roles in plants including the selective removal of regulatory proteins by the 26S proteasome. The final step in this modification is performed by ubiquitin-protein ligases (E3s) that promote Ub transfer to appropriate targets. One important family of E3s is defined by the presence of a HECT domain, an active site first found at the C-terminus of the human E3 (E6-AP). Using a consensus HECT domain as the query, we identified a family of seven HECT-containing ubiquitin-protein ligases (UPL1-UPL7) in Arabidopsis thaliana that can be grouped into four subfamilies. The UPL3 and UPL4 subfamily encodes approximately 200-kDa proteins with four Armadillo repeats similar to those in the nuclear pore protein importin-alpha, suggesting that these E3s identify their targets through binding to nuclear localization sequences. Although T-DNA disruptions of the UPL3 locus do not affect overall growth and development of Arabidopsis, the mutants show aberrant trichome morphology. Instead of developing three branches, many upl3 trichomes contain five or more branches. The upl3 trichomes also often undergo an additional round of endoreplication resulting in enlarged nuclei with ploidy levels of up to 64C. upl3 plants are hypersensitive to gibberellic acid-3 (GA3), consistent with the role of gibberellins in trichome development. The phenotype of upl3 mutants is similar to that of kaktus, a previously described set of trichome mutants with supernumerary branches. Genetic analyses confirmed that upl3 mutants and kaktus-2 are allelic with kaktus-2 plants harboring a splice-site mutation within the UPL3-transcribed region. Collectively, the data indicate that the ubiquitination of one or more activator proteins by UPL3 is necessary to repress excess branching and endoreplication of Arabidopsis trichomes.