If an enzyme name is shown in bold, there is experimental evidence for this enzymatic activity.
|Superclasses:||Biosynthesis → Hormones Biosynthesis → Plant Hormones Biosynthesis → Gibberellins and Gibberellin Precursors Biosynthesis|
|Biosynthesis → Secondary Metabolites Biosynthesis → Terpenoids Biosynthesis → Diterpenoids Biosynthesis → Gibberellins and Gibberellin Precursors Biosynthesis|
Note: This is a chimeric pathway, comprising reactions from multiple organisms, and typically will not occur in its entirety in a single organism. The taxa listed here are likely to catalyze only subsets of the reactions depicted in this pathway.
This super-pathway provides an overview of the complexity and interconnectivity of gibberellins biosynthesis. It is important to note that a given species shown here may not synthesize all the gibberellins presented. To know which gibberellins are found in a specfic species, refer to the subpathways given below.
At the time of this review, 136 fully characterized gibberellins (starting with GA1 [MacMillan68]) have been identified in more than a hundred vascular plant species, seven bacteria and seven fungi [Sponsel04, MacMillan01]. Of these gibberellins only a few have biological activity. Many of the GAs identified early in the history of the discovery of these hormones are the ones which possess the highest biological activity. These include GA1, GA3, GA4, GA5, GA6 and GA7. GA1 is the most active GA for stem elongation in Zea mays and Pisum sativum, while GA4 is the most active GA in Cucurbitaceae and in Arabidopsis thaliana. GA3 (gibberellic acid), on the other hand, has been identified in more than 40 plants and is the major GA in the fungus Fusarium fujikuroi. GA3 is used commercially to promote seed germination, stem elongation and fruit growth. In Lolium, GA5 and GA6 have been shown to enhance flowering, whilst GA1 and GA4 enhanced stem elongation [King03].
Gibberellins are diterpenes which, in higher plants, are synthesized in the plastids from glyceraldehyde-3-phosphate and pyruvate via the isopentenyl diphosphate (IPP). They all have either 19 or 20 carbon units grouped into either four or five ring systems (see C20-gibberellin skeleton and C19-gibberellin skeleton, respectively). The fifth ring is a lactone ring. GAs containing a tetracyclic ent-gibberellane structure (C20-gibberellin skeleton) are called C20-GAs, whereas GAs containing a pentacyclic 20-nor-ent-gibberellane structure (C19-gibberellin skeleton) are called C19-GAs. C20-GAs do not usually have biological activity but can be metabolized to active C19-GAs (note that not all C19-GAs are bioactive).
Gibberellins are believed to be synthesized in young tissues of the shoot and also the developing seed. It is uncertain whether young root tissues also produce gibberellins. There is also some evidence that leaves may be the source of some biosynthesis.
Active gibberellins show many physiological effects, each depending on the type of gibberellin present as well as the species of plant. Some of the physiological processes stimulated by gibberellins are: i) stimulation of stem elongation by stimulating cell division and elongation, ii) stimulation of bolting/flowering in response to long days, iii) interruption of seed dormancy in plants requiring stratification or light to induce germination, iv) stimulation of enzyme production (α-amylases) in germinating cereal grains for mobilization of seed reserves, v) induction of maleness in dioecious flowers (sex expression), vi) parthenocarpic (seedless) fruit development, and vii) delay of senescence in leaves and citrus fruits.
Certain commercial chemicals which are used to stunt growth do so in part because they block the synthesis of gibberellins. Some of these chemicals are phosphon D, AMO-1618, cycocel, ancymidol and paclobutrazol (more inhibitors of GA biosynthesis can be found in [Sponsel04]). During active growth, the plant will maintain giberellin homeostasis by metabolizing most gibberellins by rapid hydroxylation to inactive conjugates. Active gibberellin A3 is degraded much slower which helps to explain the symptoms observed in the disease bakanae caused by the rice pathogen Fusarium fujikuroi. This pathogen produces large amounts of GA3 which it secretes. Inactive conjugates might be stored or translocated via the phloem and xylem before their release (activation) at the proper time and in the proper tissue.
Subpathways: gibberellin biosynthesis II (early C-3 hydroxylation) , gibberellin biosynthesis III (early C-13 hydroxylation) , gibberellin biosynthesis I (non C-3, non C-13 hydroxylation) , gibberellin biosynthesis V , superpathway of gibberellin GA12 biosynthesis , GA12 biosynthesis , ent-kaurene biosynthesis I
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Helliwell01: Helliwell CA, Chandler PM, Poole A, Dennis ES, Peacock WJ (2001). "The CYP88A cytochrome P450, ent-kaurenoic acid oxidase, catalyzes three steps of the gibberellin biosynthesis pathway." Proc Natl Acad Sci U S A 98(4);2065-70. PMID: 11172076
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