GA20OX1

UniProt ID: Q39110
Organism: Arabidopsis thaliana
Review Status: COMPLETE
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Gene Description

GA20OX1 (GA5, AtGA20ox1; At4g25420) encodes a gibberellin 20-oxidase, a soluble 2-oxoglutarate- and Fe(II)-dependent dioxygenase that catalyses the successive oxidation of C-20 of GA12 and GA53 (via the alcohol and aldehyde) and its removal to form the C19-gibberellins GA9 and GA20, the immediate precursors of bioactive GA4 and GA1. It is one of five Arabidopsis GA20ox genes and, with GA20ox2, is the main paralog in vegetative and early reproductive development; the two act partially redundantly to promote hypocotyl and internode elongation, flowering, anther filament and silique growth. ga5 mutants are semi-dwarfs. GA20OX1 transcription is a major control point of GA levels: it is repressed by bioactive GA through DELLA-dependent feedback, induced by long days and far-red-rich light, and subject to circadian and auxin regulation.

Existing Annotations Review

GO Term Evidence Action Reason
GO:0005737 cytoplasm
ISM
GO_REF:0000122
ACCEPT
Summary: Predicted cytoplasmic localization.
Reason: GA 20-oxidases are soluble 2-oxoglutarate-dependent dioxygenases without targeting signals; the late steps of GA biosynthesis are cytosolic.
Supporting Evidence:
PMID:22533671
In higher plants, GAs are synthesized by the action of terpene cyclases, cytochrome P450 mono-oxygenases and 2-oxoglutarate-dependent dioxygenases localized, respectively, in plastids, the endomembrane system and the cytosol.
GO:0005737 cytoplasm
TAS
PMID:10330476
Feedback regulation of GA5 expression and metabolic engineer...
ACCEPT
Summary: Author statement that GA20ox is cytoplasmic.
Reason: Consistent with the cytosolic location of GA dioxygenases.
Supporting Evidence:
PMID:22533671
In higher plants, GAs are synthesized by the action of terpene cyclases, cytochrome P450 mono-oxygenases and 2-oxoglutarate-dependent dioxygenases localized, respectively, in plastids, the endomembrane system and the cytosol.
GO:0009416 response to light stimulus
IBA
GO_REF:0000033
KEEP AS NON CORE
Summary: Response to light stimulus propagated in the GA20ox clade.
Reason: GA20ox1 transcript levels respond to photoperiod and far-red light, and the gene contributes to light-regulated petiole elongation in short days; this is regulation of the enzyme rather than its core function.
Propagation Review
Root cause: NO FAILURE NON CORE
Sources checked:
PANTHER:PTN001612387 SUPPORTS TRANSFER
GA20ox clade node seeded by Arabidopsis GA20ox1/GA20ox2, whose expression is light/photoperiod regulated.
AGI_LocusCode:AT5G51810 SUPPORTS TRANSFER
GA20ox2, the most far-red-responsive GA20ox gene.
Supporting Evidence:
PMID:7604047
Expression of the GA5 gene in Ara-bidopsis leaves was enhanced after plants were transferred from short to long days; it was reduced by GA4 treatment, suggesting end-product repression in the GA biosynthetic pathway.
PMID:15923331
AtGA20ox1 also responded but to a lesser extent.
PMID:15923331
it was only in SD that a T-DNA insertional mutant of AtGA20ox1 (ga5-3) showed reduced response
GO:0009686 gibberellin biosynthetic process
IBA
GO_REF:0000033
ACCEPT
Summary: GA20ox1 oxidizes C-20 of GA12/GA53 through to the C19-GAs GA9/GA20, the immediate precursors of bioactive GA4/GA1.
Reason: Conserved role of the GA20ox clade.
Supporting Evidence:
PMID:7630935
each of which oxidized GA12 at C-20 to GA15, GA24, and the C19 compound GA9, a precursor of bioactive GAs
GO:0009686 gibberellin biosynthetic process
IDA
PMID:7630935
Isolation and expression of three gibberellin 20-oxidase cDN...
ACCEPT
Summary: GA20ox1 oxidizes C-20 of GA12/GA53 through to the C19-GAs GA9/GA20, the immediate precursors of bioactive GA4/GA1.
Reason: Recombinant enzyme converts GA12 to GA9.
Supporting Evidence:
PMID:7630935
each of which oxidized GA12 at C-20 to GA15, GA24, and the C19 compound GA9, a precursor of bioactive GAs
GO:0009686 gibberellin biosynthetic process
IEA
GO_REF:0000120
ACCEPT
Summary: GA20ox1 oxidizes C-20 of GA12/GA53 through to the C19-GAs GA9/GA20, the immediate precursors of bioactive GA4/GA1.
Reason: Consistent with experimental evidence.
Supporting Evidence:
PMID:7630935
each of which oxidized GA12 at C-20 to GA15, GA24, and the C19 compound GA9, a precursor of bioactive GAs
GO:0009686 gibberellin biosynthetic process
TAS
PMID:15923331
The involvement of gibberellin 20-oxidase genes in phytochro...
ACCEPT
Summary: GA20ox1 oxidizes C-20 of GA12/GA53 through to the C19-GAs GA9/GA20, the immediate precursors of bioactive GA4/GA1.
Reason: Consistent with experimental evidence.
Supporting Evidence:
PMID:7630935
each of which oxidized GA12 at C-20 to GA15, GA24, and the C19 compound GA9, a precursor of bioactive GAs
GO:0009739 response to gibberellin
IEP
PMID:7604047
The GA5 locus of Arabidopsis thaliana encodes a multifunctio...
KEEP AS NON CORE
Summary: GA20ox1 (GA5) transcript is down-regulated by bioactive GA (negative feedback).
Reason: Genuine transcriptional response forming part of GA homeostasis, but a regulatory property rather than the enzyme's function.
Supporting Evidence:
PMID:7604047
Expression of the GA5 gene in Ara-bidopsis leaves was enhanced after plants were transferred from short to long days; it was reduced by GA4 treatment, suggesting end-product repression in the GA biosynthetic pathway.
PMID:10330476
Negative feedback regulation of GA5 expression was demonstrated in stems of Arabidopsis by bioactive GAs but not by inactive GA.
GO:0009740 gibberellic acid mediated signaling pathway
TAS
PMID:10330476
Feedback regulation of GA5 expression and metabolic engineer...
MARK AS OVER ANNOTATED
Summary: GA20ox1 makes GA precursors; it is not a component of GA signal transduction.
Reason: Feedback regulation of GA5 by GA signaling makes GA20ox1 a target, not a component, of the pathway; the process role is GA biosynthesis.
Supporting Evidence:
PMID:10330476
Negative feedback regulation of GA5 expression was demonstrated in stems of Arabidopsis by bioactive GAs but not by inactive GA.
GO:0045544 gibberellin 20-oxidase activity
IBA
GO_REF:0000033
ACCEPT
Summary: GA 20-oxidase activity conserved in the GA20ox clade.
Reason: Experimentally supported on the target.
Supporting Evidence:
PMID:7630935
each of which oxidized GA12 at C-20 to GA15, GA24, and the C19 compound GA9, a precursor of bioactive GAs
GO:0045544 gibberellin 20-oxidase activity
IDA
PMID:7604047
The GA5 locus of Arabidopsis thaliana encodes a multifunctio...
ACCEPT
Summary: Recombinant GA5 protein converts GA53 to GA44 and GA19 to GA20; ga5 carries a nonsense mutation.
Reason: Direct enzyme assay.
Supporting Evidence:
PMID:7604047
the identity of the cDNA clone was confirmed by analyzing the capability of the fusion protein expressed in Escherichia coli to convert GA53 to GA44 and GA19 to GA20
PMID:7604047
The ga5 semidwarf mutant contains a G-->A point mutation that inserts a translational stop codon in the protein-coding sequence, thus confirming that the GA5 locus encodes GA 20-oxidase.
GO:0045544 gibberellin 20-oxidase activity
IDA
PMID:7630935
Isolation and expression of three gibberellin 20-oxidase cDN...
ACCEPT
Summary: Recombinant At2301 (GA20ox1) oxidizes GA12 to GA15, GA24 and GA9.
Reason: Core activity (RHEA:60772).
Supporting Evidence:
PMID:7630935
each of which oxidized GA12 at C-20 to GA15, GA24, and the C19 compound GA9, a precursor of bioactive GAs
GO:0045544 gibberellin 20-oxidase activity
IEA
GO_REF:0000120
ACCEPT
Summary: Rhea/ARBA mapping.
Reason: Matches experimental evidence.
Supporting Evidence:
PMID:7630935
each of which oxidized GA12 at C-20 to GA15, GA24, and the C19 compound GA9, a precursor of bioactive GAs
GO:0045544 gibberellin 20-oxidase activity
TAS
PMID:10330476
Feedback regulation of GA5 expression and metabolic engineer...
ACCEPT
Summary: Author statement on GA5 20-oxidase function.
Reason: Consistent with direct enzyme assays.
Supporting Evidence:
PMID:10330476
The gibberellin (GA) 20-oxidase encoded by the GA5 gene of Arabidopsis directs GA biosynthesis to active GAs
GO:0045544 gibberellin 20-oxidase activity
TAS
PMID:15923331
The involvement of gibberellin 20-oxidase genes in phytochro...
ACCEPT
Summary: Author statement on GA20ox function.
Reason: Consistent with direct enzyme assays.
Supporting Evidence:
PMID:7630935
each of which oxidized GA12 at C-20 to GA15, GA24, and the C19 compound GA9, a precursor of bioactive GAs
GO:0048366 leaf development
IMP
PMID:20460583
Increased leaf size: different means to an end.
KEEP AS NON CORE
Summary: GA20OX1 overexpression increases leaf size, mainly through increased cell number.
Reason: Overexpression shows that GA supply can promote leaf growth; a downstream developmental consequence of GA biosynthesis, and loss-of-function ga20ox1 ga20ox2 plants show little leaf-expansion defect, so this is non-core.
Supporting Evidence:
PMID:20460583
we selected five genes belonging to different functional classes that all positively affect leaf size when overexpressed: AVP1, GRF5, JAW, BRI1, and GA20OX1.
GO:0048575 short-day photoperiodism, flowering
IEP
PMID:7604047
The GA5 locus of Arabidopsis thaliana encodes a multifunctio...
KEEP AS NON CORE
Summary: GA5 expression increases on transfer from short to long days; GA20ox1 contributes redundantly to flowering time.
Reason: The supporting observation is transcript-level (GA5 induction on short-to-long-day transfer), which does not by itself place GA20ox1 in either short-day or long-day photoperiodic flowering. The curator's short-day term is consistent with the classical finding that GA is required for Arabidopsis flowering specifically under short days (GA-deficient ga1 mutants fail to flower in short days but flower in long days; Wilson, Heckman and Somerville 1992, Plant Physiol, not in the local publication cache), and GA20ox1/GA20ox2 act redundantly to promote flowering time (PMID:18069939, photoperiod not stated in the cached abstract). The short-day-specific ga5-3 phenotype in PMID:15923331 concerns end-of-day far-red petiole elongation, not flowering, so it does not adjudicate this term. Retained as non-core because GA20ox1 supplies GA precursors and the flowering role is a downstream consequence of GA biosynthesis.
Supporting Evidence:
PMID:7604047
Expression of the GA5 gene in Ara-bidopsis leaves was enhanced after plants were transferred from short to long days; it was reduced by GA4 treatment, suggesting end-product repression in the GA biosynthetic pathway.
PMID:18069939
AtGA20ox1 and AtGA20ox2 act redundantly to promote hypocotyl and internode elongation, flowering time, elongation of anther filaments, the number of seeds that develop per silique and elongation of siliques

Core Functions

Cytosolic 2-oxoglutarate/Fe(II)-dependent dioxygenase that oxidizes C-20 of GA12 (and GA53) in successive steps via GA15 and GA24 to the C19-gibberellin GA9 (and GA20), with loss of C-20 as CO2 (RHEA:60772); a major rate-limiting, feedback-regulated step in bioactive GA formation, acting partially redundantly with GA20ox2.

Cellular Locations:
Supporting Evidence:
  • PMID:7630935
    each of which oxidized GA12 at C-20 to GA15, GA24, and the C19 compound GA9, a precursor of bioactive GAs
  • PMID:7604047
    the identity of the cDNA clone was confirmed by analyzing the capability of the fusion protein expressed in Escherichia coli to convert GA53 to GA44 and GA19 to GA20
  • PMID:7604047
    The ga5 semidwarf mutant contains a G-->A point mutation that inserts a translational stop codon in the protein-coding sequence, thus confirming that the GA5 locus encodes GA 20-oxidase.
  • PMID:18069939
    AtGA20ox1 and AtGA20ox2 act redundantly to promote hypocotyl and internode elongation, flowering time, elongation of anther filaments, the number of seeds that develop per silique and elongation of siliques

References

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Suggested Questions for Experts

Q: What fraction of GA20OX1 product formed in vivo is GA9 versus the 13-hydroxylated GA20, and does the relative use of the non-13-hydroxylation and early-13-hydroxylation routes vary by tissue?

Suggested Experiments

Experiment: Quantify GA12/GA53-derived intermediates by isotope-dilution LC-MS in ga20ox1 single and ga20ox1 ga20ox2 double mutant organs to assign the in vivo contribution of GA20ox1 to each branch.

πŸ“š Additional Documentation

Notes

(GA20OX1-notes.md)

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