Role Of Gibberellins Research Articles

Overexpression of the gibberellin 20-oxidase gene from Torenia fournieri resulted in modified trichome formation and terpenoid metabolities of Artemisia annua L.

Gibberellins (GAs) are diterpenoid hormones, control various physiological developments in plants. The role of gibberellins on morphology and secondary metabolite production was examined in Artemisia annua, a medicinal plant that has been acknowledged as a source of artemisinin, an antimalarial compound. Subsequently, the GA20ox gene from Torenia fournieri (TfGA20ox2) was transferred to A. annua by Agrobacterium-mediated transformation. Compared with wild type plants, all nine transgenic plants showed significantly higher plant heights and artemisinin contents. The highest artemisinin content and yield in TfGA20ox2-overexpressing plants was around two-fold higher than wild type. Moreover, transgenic plants had higher numbers of branches (52.4%) and greater branch lengths (60–203%), but smaller leaf size (77.6%). Interestingly, relative to wild type the number and size of glandular trichomes in transgenic leaves was about 30 and 35% higher, respectively. From GC–MS analysis, the proportion of diterpenes in transgenic plant extracts was 1.5-fold lower than those noticed in wild type, while the proportion of sesquiterpenes was increased about 1.6 times when compared to wild type. However, the content proportion of monoterpenes showed a slightly increase, whereas the level of triterpenes showed no variation. In addition, two monoterpenes (eucalyptol and borneol), four sesquiterpenes (α-caryophyllene, β-guaiene, δ-cadinene and β-cubebene) and one triterpenes (isomultiflorenone) were detected only in transgenic extract, whereas d-α-tocopherol, a diterpenoid compound was found only in wild type but not transgenic plant. These results suggested that gibberellins play a significant role in regards to morphology, trichome formation and terpenoid metabolite production in A. annua.

Rice Root-knot Nematode Down Regulates GA-Gene to Establish Infection

Experiments were conducted to observe the role of gibberellin (GA) in the response of rice to infection by rice root-knot nematode, Meloidogyne graminicola. Gibberellin might have a negative effect on defence in Nipponbare against M. graminicola. On the other hand, an inhibitory effect to M. graminicola in Taichung 65 (wild type) was found when GA was applied exogenously. Galling was significantly and reproducibly lower in mutants, whether it was GA insensitive or GA deficient, than in Taichung 65. There were even significantly lower galls in these GA deficient mutants after GA treatment than in non-treated mutant plants. The expression of GA signaling gene Osgid1-7 was significantly higher only in whole root at 2 dpi (days post inoculation) but not in leaves or root tips. Even this gene was not differentially expressed in root and leaf tissues at later stage (6 dpi days post inoculation) of infection. The nematode inhibited biosynthesis of OsGA20ox in leaves at 2 dpi and, in both of leaves and roots at 6 dpi. The GA response gene AK073385 was also less expressed in root tip at 6 dpi (days post inoculation). These might help M. graminicola to continue their infection in Taichung 65.

The role of gibberellins in improving the resistance of tebuconazole-coated maize seeds to chilling stress by microencapsulation.

Chilling stress during germination often causes severe injury. In the present study, maize seed germination and shoot growth under chilling stress were negatively correlated with the dose of tebuconazole in an exponential manner as predicted by the model Y = A + B × e(−x/k). Microencapsulation was an effective means of eliminating potential phytotoxic risk. The gibberellins (GAs) contents were higher after microencapsulation treatment than after conventional treatment when the dose of tebuconazole was higher than 0.12 g AI (active ingredient) kg−1 seed. Further analysis indicated that microencapsulation can stimulate ent-kaurene oxidase (KO) activity to some extent, whereas GA 3-oxidase (GA3ox) and GA 2-oxidase (GA2ox) activities remained similar to those in the control. Genes encoding GA metabolic enzymes exhibited different expression patterns. Transcript levels of ZmKO1 increased in the microcapsule treatments compared to the control. Even when incorporated into microcapsules, tebuconazole led to the upregulation of ZmGA3ox1 at doses of less than 0.12 g AI kg−1 seed and to the upregulation of ZmGA3ox2 when the dose was higher than 0.12 g AI kg−1 seed. With increasing doses of microencapsulated tebuconazole, the transcript levels of ZmGA2ox4, ZmGA2ox5 and ZmGA2ox6 exhibited upward trends, whereas the transcript levels of ZmGA2ox7 exhibited a downward trend.

The role of gibberellins and auxin on the tomato cell layers in pericarp via the expression of ARFs regulated by miRNAs in fruit set

Phytohormones, such as auxin (IAA) and gibberellin (GA), are known to be essential for fruit development. We utilized GA-deficient (gib-3) and diageotropica (dgt) tomato mutants to elucidate the effects of single hormones in the pericarp. The application of IAA or GA, respectively, to gib-3 or dgt single mutants induced a significant morphological difference in the fruit set. We found that IAA application induced cell division in the gib-3 pericarp and that GA application did not increase the cell layers in the dgt pericarp. In molecular studies, the expression levels of SlARF6, SlARF8, SlARF10 and SlARF16 were downregulated by IAA application, whereas the expression of their regulators miRNA160 and miRNA167 was upregulated by IAA application in gib-3 plants. Furthermore, the expression levels of SlARF6, SlARF8, SlARF10 and SlARF16 were upregulated by GA application, whereas the expression levels of miRNAs were reduced in the dgt mutant. These results imply that the expression levels of SlARF6, SlARF8, SlARF10 and SlARF16 were negatively correlated with the number of cell layers in the pericarp during fruit set. To further support this hypothesis, 35s:mSlARF10 transgenic plants resistant to SlmiR160 cleavage of SlARF10 mRNA were used to investigate the cell layers in fruit. These results revealed that mSlARF10 overexpression indeed resulted in fewer cell layers than in wild type fruit. Together, our data suggest that GA- and IAA-mediated miRNAs and their target ARFs influence the formation of pericarp cell layers during fruit set development.

Roles of Gibberellins and Abscisic Acid in Regulating Germination of Suaeda salsa Dimorphic Seeds Under Salt Stress.

Seed heteromorphism observed in many halophytes is an adaptive phenomenon toward high salinity. However, the relationship between heteromorphic seed germination and germination-related hormones under salt stress remains elusive. To gain an insight into this relationship, the roles of gibberellins (GAs) and abscisic acid (ABA) in regulating germination of Suaeda salsa dimorphic brown and black seeds under salinity were elucidated by studying the kinetics of the two hormones during germination of the two seed types with or without salinity treatment. Morphological analysis suggested that brown and black are in different development stage. The content of ABA was higher in dry brown than in black seeds, which gradually decreased after imbibition in water and salt solutions. Salt stress induced ABA accumulation in both germinating seed types, with higher induction effect on black than brown seeds. Black seeds showed lower germination percentage than brown seeds under both water and salt stress, which might be attributed to their higher ABA sensitivity rather than the difference in ABA content between black and brown seeds. Bioactive GA4 and its biosynthetic precursors showed higher levels in brown than in black seeds, whereas deactivated GAs showed higher content in black than brown seeds in dry or in germinating water or salt solutions. High salinity inhibited seed germination through decreasing the levels of GA4 in both seeds, and the inhibited effect of salt stress on GA4 level of black seeds was more profound than that of brown seeds. Taken together higher GA4 content, and lower ABA sensitivity contributed to the higher germination percentage of brown seeds than black seeds in water and salinity; increased ABA content and sensitivity, and decreased GA4 content by salinity were more profound in black than brown seeds, which contributed to lower germination of black seeds than brown seeds in salinity. The differential regulation of ABA and GA homeostases by salt stress in dimorphic seeds might provide a strategy for S. salsa plants to survive adverse environmental conditions.

Exogenous gibberellin altered morphology, anatomic and transcriptional regulatory networks of hormones in carrot root and shoot.

BackgroundGibberellins stimulate cell elongation and expansion during plant growth and development. Carrot is a root plant with great value and undergoes obvious alteration in organ size over the period of plant growth. However, the roles of gibberellins in carrot remain unclear.ResultsTo investigate the effects of gibberelliins on the growth of carrot, we treated carrot plants with gibberellic acid 3 (GA3) or paclobutrazol (a gibberellin inhibitor). The results found that GA3 dramatically reduced the root growth but stimulated the shoot growth of carrot. It also significantly promoted xylem development in the tuberous root of carrot. In addition, transcript levels of genes related to gibberellins, auxin, cytokinins, abscisic acid and brassinolides were altered in response to increased or reduced gibberellins.ConclusionsThe inhibited tuberous root growth but enhanced shoot growth in plants treated with GA3 can be principally attributed to the changes in the xylem development of carrot roots. Negative feedback regulation mechanism of gibberellin biosynthesis also occurred in response to altered gibberellin accumulation. Gibberellins may interact with other hormones to regulate carrot plant growth through crosstalk mechanisms. This study provided novel insights into the functions of gibberellins in the growth and development of carrot.

The role of gibberellins in the mitigation of chilling injury in cherry tomato (Solanum lycopersicum L.) fruit

Tomato fruit is cold-sensitive and susceptible to chilling injury during cold storage. Bioactive gibberellins (GAs), which play an important role in regulating various physiological processes, have been reported to be involved in stress responses of plant. In this study, we investigated the effect of GA on chilling injury of harvested cherry tomato fruit. We found that low temperature significantly inhibited increase of endogenous levels of gibberellic acid (GA3) in fruit. This effect was associated with lower expression of key GA metabolic genes, GA3ox1, GA20ox1 and GA2ox1. GA3 treatment reduced the chilling injury index, whereas treatment with paclobutrazol (PAC), a GA biosynthesis inhibitor, exacerbated chilling injury. Compared with the control, GA3 treatment decreased electrolyte leakage and malondialdehyde content, increased proline content, and improved antioxidant enzyme activities. Treatment with PAC caused the opposite effects. These results suggest that endogenous GA levels were positively associated with lipid peroxidation and oxidative stress in chilled fruit. GA appears to play an important role in cold response regulation in fruit.

Understanding flowers and flowering: an integrated approach

INTRODUCTION 1. The Evolution of Flowers 2. Historical Interpretations of Flower Induction and Flower Development THE MOLECULAR MECHANISMS OF FLOWERING: INDUCTION AND DEVELOPMENT 3. Flower induction in Arabidopsis thaliana 4. The autonomous pathways for floral inhibition and induction 5. The photoperiodic pathway of floral induction 6. The vernalization pathway of floral induction and the role of gibberellin 7. Integrating the Arabidopsis thaliana flower induction pathways and assessing the extent to which the model is ubiquitous 8. Changes at the shoot apical meristem in response to floral induction 9. Development of the floral organs 10. The ABC model in evolution 11. Development of gametes 12. Preventing self-fertilisation POLLINATION SUCCESS: MOLECULAR AND ECOLOGICAL INTERACTIONS 13. Why are flowers different? Pollination syndromes - the theory 14. Changing floral shape and structure 15. Colouring the flower 16. Enhancing flower colour 17. Are flowers under selective pressure to increase pollinator attention? 18. Do pollinators discriminate between different floral forms? 19. Pollination syndromes - the evidence Epilogue Bibliography

Role of gibberellins during arbuscular mycorrhizal formation in tomato: new insights revealed by endogenous quantification and genetic analysis of their metabolism in mycorrhizal roots.

Gibberellins (GAs) are key regulators of plant growth and development and recent studies suggest also a role during arbuscular mycorrhizal (AM) formation. Here, complementary approaches have been used to obtain a clearer picture that correlates AM fungal development inside roots with GA metabolism. An extensive analysis of genes associated with GA metabolism as well as a quantification of GA content in roots was made. Application of GA3 and its biosynthesis inhibitor prohexadione calcium (PrCa) combined with a GA-constitutive response mutant (procera) were used to determine whether fungal colonization is altered by the level of these hormones or by changes in the GA-signaling pathway. The increased levels of specific GAs from the 13-hydroxylation pathway in mycorrhizal roots correlate closely with the increased expression of genes coding enzymes from the GA biosynthetic trail. The imbalance of GAs in tomato roots caused by exogenous applications of GA3 or PrCa affects arbuscules in both negative and positive ways, respectively. In addition, procera plants were adversely affected by the mycorrhization process. Our findings demonstrate that an imbalance in favor of an increased amount of GAs negatively affects the frequency of mycorrhization and particularly the arbuscular abundance in tomato mycorrhizal roots and the results point out that AM formation is associated with a change in the 13-hydroxylation pathway of GAs.

Roles of Gibberellin and Auxin in Promoting Seed Germination and Seedling Vigor in Pinus massoniana

Roles of Gibberellin and Auxin in Promoting Seed Germination and Seedling Vigor in Pinus massoniana

Removal of DELLA repression promotes leaf senescence in Arabidopsis

Leaf senescence is an integrated response of leaf cells to developmental age and various internal and environmental signals. However, the role of gibberellins (GA) in leaf senescence is not clear. In the current study, we investigated the effect of DELLA on leaf senescence. Compared with the wild type (WT), leaf senescence occurred earlier in the mutant ga1-3 gai-t6 rga-t2 rgl1-1 rgl2-1 (abbreviated as Q-DELLA/ga1-3) whose DELLA repression was removed, whereas leaf senescence was retarded in the mutant ga1-3 whose GA biosynthesis was blocked and whose DELLA proteins accumulated abnormally. During leaf senescence, SAG12 and SAG29 were upregulated in Q-DELLA/ga1-3 and downregulated in ga1-3 plants. The Q-DELLA/ga1-3 senescent leaves contained more sugar but less chlorophyll and fatty acids (FAs) than those of ga1-3 and WT. Both absolute and relative contents of C18:3 in Q-DELLA/ga1-3 senescent leaves were lower compared with those of the WT and ga1-3 leaves. The genes regulating FA β-oxidation in Q-DELLA/ga1-3, such as KAT2, LACS6, LACS7, ACX1, ACX2 and MAP2, were significantly upregulated. The removal of DELLA repression highly upregulated certain genes on various hormone pathways, suggesting that GA signaling acts upstream of the jasmonic acid, salicylic acid, and ethylene pathways in regulating leaf senescence.

The Role of Gibberellin in the CBF1-Mediated Stress-Response Pathway

Constitutive expression of C-repeat/dehydration-responsive element binding factor genes confers drought, salt, and freezing tolerance in transgenic plants but causes dwarfism under normal growth conditions. Evidence has emerged that gibberellin (GA) plays a major role in these growth-regulatory mechanisms. However, the role and benefit of reducing the active GA level in abiotic stress responses are largely unknown. Using microarray analysis, we identified a subgroup of GA-regulated genes by comparing the transcriptome of GA over-accumulation and deficiency, CBF1 constitutive expression, and wild-type tobacco plants (Nicotiana tabacum). The results revealed that the geranylgeranyl diphosphate synthase is a regulatory site integrating environmental cues and is endogenous in the regulation of GA level and subsequent plant growth. Low levels of GA promoted the accumulation and gene activation of osmotic adjustment-related substances and enhanced the water-retention capability of leaves. We found that GA was associated with SA/JA balance in the CBF1-mediated stress response, but inhibition of SA signaling is the principal regulator in this pathway. These results demonstrate that, in addition to regulating growth, GA has a much more complex role, and SA may be another important player in the CBF1-mediated stress-response pathway.

Plant growth enhancement and associated physiological responses are coregulated by ethylene and gibberellin in response to harpin protein Hpa1

The harpin protein Hpa1 produced by the bacterial blight pathogen of rice induces several growth-promoting responses in plants, activating the ethylene signaling pathway, increasing photosynthesis rates and EXPANSIN (EXP) gene expression levels, and thereby enhancing the vegetative growth. This study was attempted to analyze any mechanistic connections among the above and the role of gibberellin in these responses. Hpa1-induced growth enhancement was evaluated in Arabidopsis, tomato, and rice. And growth-promoting responses were determined mainly as an increase of chlorophyll a/b ratio, which indicates a potential elevation of photosynthesis rates, and enhancements of photosynthesis and EXP expression in the three plant species. In Arabidopsis, Hpa1-induced growth-promoting responses were partially compromised by a defect in ethylene perception or gibberellin biosynthesis. In tomato and rice, compromises of Hpa1-induced growth-promoting responses were caused by a pharmacological treatment with an ethylene perception inhibitor or a gibberellin biosynthesis inhibitor. In the three plant species, moreover, Hpa1-induced growth-promoting responses were significantly impaired, but not totally eliminated, by abolishing ethylene perception or gibberellin synthesis. However, simultaneous nullifications in both ethylene perception and gibberellin biosynthesis almost canceled the full effects of Hpa1 on plant growth, photosynthesis, and EXP2 expression. Theses results suggest that ethylene and gibberellin coregulate Hpa1-induced plant growth enhancement and associated physiological and molecular responses.Electronic supplementary materialThe online version of this article (doi:10.1007/s00425-013-2013-y) contains supplementary material, which is available to authorized users.

Studies on the role of gibberellins in the regulation of spermatogenesis in Chara vulgaris L.

Antheridia from isolated nodes of Chara vulgaris, developing in the presence of either AMO-1618 or GA<sub>3</sub>, were studied. AMO-1618 which lowers the level of endogenous gibberellins causes a significant, proportional to the concentration, reduction in: 1) the number of antheridial filaments formed in antheridia, 2) spermatid number within a filament, as a result of eliminating one mitotic division at the first stage of spermatogenesis. Exogenous GA<sub>3</sub> at the concentration 10<sup>-5</sup> M evokes opposite effect i.e. increase in the number of antheridial filaments and increase in the number of spermatids within filament. Total number of spermatids within an antheridium decreases under the influence of 10<sup>-4</sup>M AMO-1618 three times in comparison with the control, whereas it increases twice following 10<sup>-5</sup>M GA<sub>3</sub> treatment. It has been suggested that the normal course of spermatogenesis requires precisely determined level of endogenous gibberellins.

Role of gibberellins and cytokinins in regulation of germination during development and ripening of Triticale caryopses

The germination of caryopses of M-T3 <em>Triticale</em> generation, which were freshly harvested in different growth and developmental phases has been studied. A significant influence of the abscisic acid (ABA) accumulation on the increment of number of germinating caryopses has been found. Already in the first phase af the embryogenesis considerable stimulating effects of kinetin and gibberellin-A<sub>3</sub> (GA<sub>3</sub>) on the germination of embryos which were isolated from freshly collected grains have been shown. When both stimulators were used together marked synergetic reaction occurred. It has been also determined that in the initial period of embryogensis premature germination occurs, to a higher extent, under the action of cytokinins than! Whether in the further phases of the caryopse development, when embryo develop mainly through the cell elongation, mostly gibberellins seem to be responsible for the activation of germination processes. The more mature were seeds the quicker germinated whole caryopses and embryos isolated from them at different ripeness, after 3-month storage. The highest stimulation of germination by phytohormones has been found for the most mature caryopses. The action of gibberellic acid has been particulary strong.

Plant hormones in arbuscular mycorrhizal symbioses: an emerging role for gibberellins.

Arbuscular mycorrhizal symbioses are important for nutrient acquisition in >80 % of terrestrial plants. Recently there have been major breakthroughs in understanding the signals that regulate colonization by the fungus, but the roles of the known plant hormones are still emerging. Here our understanding of the roles of abscisic acid, ethylene, auxin, strigolactones, salicylic acid and jasmonic acid is discussed, and the roles of gibberellins and brassinosteroids examined. Pea mutants deficient in gibberellins, DELLA proteins and brassinosteroids are used to determine whether fungal colonization is altered by the level of these hormones or signalling compounds. Expression of genes activated during mycorrhizal colonization is also monitored. Arbuscular mycorrhizal colonization of pea roots is substantially increased in gibberellin-deficient na-1 mutants compared with wild-type plants. This is reversed by application of GA3. Mutant la cry-s, which lacks gibberellin signalling DELLA proteins, shows reduced colonization. These changes were parallelled by changes in the expression of genes associated with mycorrhizal colonization. The brassinosteroid-deficient lkb mutant showed no change in colonization. Biologically active gibberellins suppress arbuscule formation in pea roots, and DELLA proteins are essential for this response, indicating that this role occurs within the root cells.

Gibberellins regulate the transcription of the continuous flowering regulator, RoKSN, a rose TFL1 homologue

The role of gibberellins (GAs) during floral induction has been widely studied in the annual plant Arabidopsis thaliana. Less is known about this control in perennials. It is thought that GA is a major regulator of flowering in rose. In spring, low GA content may be necessary for floral initiation. GA inhibited flowering in once-flowering roses, whereas GA did not block blooming in continuous-flowering roses. Recently, RoKSN, a homologue of TFL1, was shown to control continuous flowering. The loss of RoKSN function led to continuous flowering behaviour. The objective of this study was to understand the molecular control of flowering by GA and the involvement of RoKSN in this inhibition. In once-flowering rose, the exogenous application of GA3 in spring inhibited floral initiation. Application of GA3 during a short period of 1 month, corresponding to the floral transition, was sufficient to inhibit flowering. At the molecular level, RoKSN transcripts were accumulated after GA3 treatment. In spring, this accumulation is correlated with floral inhibition. Other floral genes such as RoFT, RoSOC1, and RoAP1 were repressed in a RoKSN-dependent pathway, whereas RoLFY and RoFD repression was RoKSN independent. The RoKSN promoter contained GA-responsive cis-elements, whose deletion suppressed the response to GA in a heterologous system. In summer, once-flowering roses did not flower even after exogenous application of a GA synthesis inhibitor that failed to repress RoKSN. A model is presented for the GA inhibition of flowering in spring mediated by the induction of RoKSN. In summer, factors other than GA may control RoKSN.

Tomato SlDREB gene restricts leaf expansion and internode elongation by downregulating key genes for gibberellin biosynthesis.

Plants have evolved and adapted to different environments. Dwarfism is an adaptive trait of plants that helps them avoid high-energy costs under unfavourable conditions. The role of gibberellin (GA) in plant development has been well established. Several plant dehydration-responsive element-binding proteins (DREBs) have been identified and reported to be induced under abiotic and biotic stress conditions. A tomato DREB gene named SlDREB, which is a transcription factor and was cloned from cultivated tomato M82, was found to play a negative role in tomato plant architecture and enhances drought tolerance. Tissue expression profiles indicated that SlDREB was expressed mainly in the stem and leaf and could be induced by abscisic acid (ABA) but suppressed by GA and ethylene. SlDREB altered plant morphology by restricting leaf expansion and internode elongation when overexpressed, and the resulting dwarfism of tomato plants could be recovered by application of exogenous gibberellic acid (GA3). Transcriptional analysis of transgenic plants revealed that overexpression of SlDREB caused the dwarf phenotype by downregulating key genes involved in GA biosynthesis such as ent-copalyl diphosphate synthase (SlCPS) and GA 20-oxidases (SlGA20ox1, -2, and -4), thereby decreasing endogenous GA levels in transgenic plants. A yeast activity assay demonstrated that SlDREB specifically bound to dehydration-responsive element/C-repeat (DRE/CRT) elements of the SlCPS promoter region. Taken together, these data demonstrated that SlDREB can downregulate the expression of key genes required for GA biosynthesis and that it acts as a positive regulator in drought stress responses by restricting leaf expansion and internode elongation.

Seasonal differences in the capacity of perennial ryegrass to respond to gibberellin explained

Perennial ryegrass plants (Lolium perenne) were taken from an established field at two different stages in the season (mid-winter and again at mid-summer). They were then grown in a controlled environment to both "lock in" their contrasting developmental states and to look at the role of nitrogen supply, temperature, and developmental state separately to evaluate the potential of plants to respond to exogenous application of gibberellin. Responses to exogenous gibberellin (gibberellic acid, GA) were significant but were far smaller in summer-derived than winter-derived plants. The major difference in response to GA (compared with controls) between winter-derived and summerderived plants suggests that seasonal changes in plant developmental state have a major effect in the field on the capacity for the plants to respond to exogenous GA application. This effect is greater than that of temperature and N availability. This raises new prospects for making sustained increases in plant growth, but only if the fundamental mechanisms by which plants control their responses to environmental signals (e.g., temperature and soil N status) can be understood. The role of gibberellins (endogenous as well as externally applied) in changes in plant growth strategy presents a new challenge for forage plant science.

Gibberellin Metabolism and Transport During Germination and Young Seedling Growth of Pea (Pisum sativum L.)

The role of gibberellins (GAs) during germination and early seedling growth is examined by following the metabolism and transport of radiolabeled GAs in cotyledon, shoot, and root tissues of pea (Pisum sativum L.) using an aseptic culture system. Mature pea seeds have significant endogenous GA20 levels that fall during germination and early seedling growth, a period when the seedling develops the capacity to transport GA20 from the cotyledon to the shoot and root of the seedling. Even though cotyledons at 0–2 days after imbibition have appreciable amounts of GA20, the cotyledons retain the ability to metabolize labeled GA19 to GA20 and express significant levels of PsGA20ox2 message (which encodes a GA biosynthesis enzyme, GA 20-oxidase). The large pool of cotyledonary GA20 likely provides substrate for GA1 synthesis in the cotyledons during germination, as well as for shoots and roots during early seedling growth. The shoots and roots express GA metabolism genes (PsGA3ox genes which encode GA 3-oxidases for synthesis of bioactive GA1, and PsGA2ox genes which encode GA 2-oxidases for deactivation of GAs to GA29 and GA8), and they develop the capacity to metabolize GAs as necessary for seedling establishment. Auxins also show an interesting pattern during early seedling growth, with higher levels of 4-chloro-indole-3-acetic acid (4-Cl-IAA) in mature seeds and higher levels of indole-3-acetic acid (IAA) in young root and shoot tissues. This suggests a changing role for auxins during early seedling development.