The Related Research of the Genes of Gibberellin Receptor（GID1）in Plant

Jianming WU，Xing HUANG，Lihang QlU，Huanzhong SONG，Yangrui Ll，*，Litao YANG，Ll Qiang，Liu YANG

1.Sugarcane Research Center，Chinese Academy of Agricultural Sciences，Nanning 530007，China；

2.Sugarcane Research Institute，Guangxi Academy of Agricultural Sciences，Nanning 530007，China；

3.Key Laboratory of Sugarcane Biotechnology and Genetic Improvement（Guangxi），Ministry of Agriculture，Nanning 530007 China；

4.Guangxi Crop Genetic Improvement and Biotechnology Lab，Nanning 530007，China

The Related Research of the Genes of Gibberellin Receptor（GID1）in Plant

Jianming WU1，2，3，Xing HUANG1，2，3，Lihang QlU1，2，3，Huanzhong SONG4，Yangrui Ll1，2，3，4*，Litao YANG4，Ll Qiang4，Liu YANG4

1.Sugarcane Research Center，Chinese Academy of Agricultural Sciences，Nanning 530007，China；

2.Sugarcane Research Institute，Guangxi Academy of Agricultural Sciences，Nanning 530007，China；

3.Key Laboratory of Sugarcane Biotechnology and Genetic Improvement（Guangxi），Ministry of Agriculture，Nanning 530007 China；

4.Guangxi Crop Genetic Improvement and Biotechnology Lab，Nanning 530007，China

The hormone gibberellin（GA）plays an important role in modulating diverse processes throughout plant development.Gibberellin（GA）perception is mediated by GID1（GA-INSENSITIVE DWARF1），a receptor that shows similarity to hormone-sensitive lipases.It has been postulated that plants have two types of GA receptors，including soluble and membrane-bound forms.In recent years，significant advances have been made in the research of Gibberellin Receptor（GID1）.This article highlights recent advances in the molecular structure of Gibberellin Receptor（GID1），Molecular Interactions between the Gibberellin Receptor（GID1）with DELLA，Cloning and expression of GA receptors（GID1），function identification of GA receptor gene（GID1）.These discoveries open new prospects for the understanding mechanism of GA receptors（GID1）in plants.

Gibberellin Receptor；DELLA；GID1；Gibberellin

G ibberellins （GAs）are plant hormones that regulate growth and influence various developmentalprocesses，including stem elongation，germination，dormancy，flowering，sex expression，enzyme induction，and leaf and fruit senescence［1-3］.The complex pathways of GA metabolism have been elucidated，and several factors that influence GA metabolism have been identified［4］.Impressive advances have been made in elucidating the GA pathway with the cloning and characterization of genes encoding most GA biosynthesis and catabolism enzymes，GA receptors（GIBBERELLIN INSENSITIVE DWARF1，GID1）and early GA signaling components.Recently，the GA receptor（GID1）was identified by a combination ofbiochemicaland genetic techniques［5-6］.In 2008，the structure of GA receptor have been elucidated［7-8］Because of the identification of the GA receptor，the molecular mechanisms of GA receptor and signal transduction are much better understood.The research reviewed the history of the related research of Gibberellin Receptor genes in plant，the biochemical and physiological characteristics of the GA receptor，and interactions between the Gibberellin Receptor with DELLA，and cloning and expression of GA receptors，its function identification in plant.

The Discovery of GA Receptors and ldentified Molecular Structure

The biochemical pathway of GA biosynthesis and catabolism in plants are well defined，and genes encoding most enzymes in this pathway have been identified［9］.Recent genetic，biochemical，and structural studies have elucidated the molecular mechanism of GA perception and initial steps in GA signaling in plants［10］.

In the early stages，some researchers show that GA perception occurs by a mechanism similar to hydrophobic steroid hormones in mammalian cells［10］.Johri and Varner［11］showed thatnucleiisolated from shoots of light-grow dwarf peas in the presence of gibberellic acid （10-8M）show a higher rate of DNA-dependent RNA synthesis than the untreated nuclei，RNA synthesized by GA-treated nuclei has a different nearest-neighbor frequency and also has a higher average molecular weight than the RNA synthesized by untreated nuclei.Later，Sechley and Srivastava［12］Reported addition of［3H］GA4-binding cytosolic protein to nuclei increased the overall transcription but did not affect the GA4-enhanced transcription，Receptor-binding assay indicated that these nuclei contain ［3H］GA4-binding sites which are heat-labile and bound to nuclear matrix.Further，Witham and Hendry［13］show that Computer modeling and molecularmechanics performed on the intercalation complexes of selected gibberellins or biosynthetic precursors with DNA dinucleotides revealed that under appropriate condi-tions the ligands insert（intercalate）between the base-paired double-stranded dinucleotide，5'-dTdA-3'.

On the other hand，Some scientists hold opposite viewpoint，Stoddart et al.［14］initial studies on the stabilityl and properties of the 2KP-［3H］GA1 complex from lettuce hypocotyl sections.Later，a number of GBPs have been proposed as GA receptor candidates［15-17］.Further studies have revealed that this GBP in the partially purified fraction fulfills all four of the GA receptor criteria［18-19］.The biochemical search for GA receptors has gone on for a long time，and through these experiments it has been postulated that there are two types of GA receptors，including soluble and membranebound forms［10］.

Gibberellins（GAs）has been postulated that plants have both membrane-bound and soluble GA receptors；however，no GA receptors have yet been identified.The GA receptor GIBBERELLIN INSENSITIVE DWARF1（GID1）was first identified by studies of GA-insensitive dwarf rice mutants［5］.This exciting discovery not only provides a major new piece to the puzzle of the GA signal transduction pathway but also raises new questions and calls into question some long-held assumptions concerning the topology of the pathway［20］.Recently，a soluble GA receptor，GID1，was identified in rice and in several other plant species［6，21-24］.

Although，the gibberellin receptors have been identified，the precise manner in which GID1 discriminates and becomes activated by bioactive gibberellins for specific binding to DELLA proteins remains unclear.Recently，Murase et al.［7］present the crystal structure of a ternary complex of Arabidopsis thaliana GID1A，a bioactive gibberellin and the amino-terminal DELLA domain of GAI.In this complex，GID1A occludes gibberellin in a deep binding pocket covered by its N-terminal helical switch region，which in turn interacts with the DELLA domain containing DELLA，VHYNP and LExLE motifs.Shimada et al.［8］was analysed the crystal structure of Oryza sativa GID1 show that the overall structure of both complexes shows an α/β-hydrolase fold similar to that of HSLs except for an amino-terminal lid. The GA-binding pocket corresponds to the substrate-binding site of HSLs. These results establish a structural model of a plant hormone receptor that is distinct from the mechanism of the hormone perception and effector recognition of the known auxin receptors. Furthermore， comparison of amino acid sequences of GID1 proteins among various plant species revealed that this loop region is highly diverse relative to other GID1 regions［22］.

Molecular Interactions between the Gibberellin Receptor with DELLA

DELLA proteins play a key role in growth regulatory signalling pathways in plants.GA acts，in part through destabilisation of a family of related proteins known as DELLA proteins. There are 5 known DELLA proteins in Arabidopsis，three of these are involved in GA response:GIBBERELLIC ACID INSENSTIVE （GAI），REPRESSOR OF GA1-3 （RGA）and RGA-LIKE1 （RGL1）［25-26］，which display overlapping but also some distinct functions in repressing GA responses［27-30］.But monocots are different with dicots.For example，rice and barley，only contain a single DELLA gene SLENDER1 （SLR1）in rice and SLENDER in barley［31-33］.Some studies have revealed that DELLA proteins have broad interactions with other hormonal and environmental signaling，playing important roles in many aspects of plant growth，development and adaptation to environmental stresses［34-36］.

Early results showed that angiosperm DELLAs contain in their N termini two highly conserved domains（Ⅰ and Ⅱ）that are necessary for GID1-DELLA interactions［37-38］.Functional DELLA orthologs have been identified in many angiosperms［39］.DELLA proteins are GA-signalling repressors that block GA-induced development［40］.The first evidence that the GID1 receptor is directly involved in the DELLA-mediated GA signaling system was obtained from a GA-dependent interaction between GID1 and SLR1 in a yeast two-hybrid（Y2H）assay［5］.So，it has been determined that GID1 directly interacts with DELLA proteins in a GA-dependent manner［5，21，6，41］.Griffiths et al.［21］study shows that loss of the three Arabidopsis GID1 receptors results in GA insensitivity and that the N-terminal DELLA and VHYNP domains of the DELLA protein RGA are required for GID1 interactions in Arabidopsis.While Willige et al.［41］concluded that the VHYNP domain is not essential for the interaction between GAI and At GID1a.In rice （Oryza sativa）and Arabidopsis thaliana，gibberellin（GA）signaling is mediated by GIBBERELLIN-INSENSITIVE DWARF1（GID1）and DELLA proteins in collaboration with a GA-specific F-box protein［22］.

Another model was further refined by Stoddart（1979）showing that the GRAS domain of the rice DELLA（SLR1）also interacts with GID1 to stabilize this protein complex，but this interaction only occurs after the aminoterminal DELLA/VHYNP region has bound to GID1.This model is supported by anotherstudy showing that GID1 and DELLA are co-immunopreciptated with SLY1 from protein extracts prepared from transgenic Arabidopsis overexpressing both FLAG-tagged SLY1 and HA-tagged GID1［14］. Furthermore，A recent study reveals that a loop region in OsGID1 plays a critical role in the GA-dependent interaction of GID1 with DELLA，this model suggests thatthe GA-independent binding of OsGID1P99A and AtGID1B to DELLA is caused by an altered conformation of the N-Ex so that it resembles a partially closed lid even without GA binding［42］.Some reports show that overexpression of GID1 rescues the dwarf phenotype of the sly1 and gid2 mutants withoutreducing DELLA levels，suggesting that GID1-DELLA interaction can inhibit DELLA function without protein degradation［43-44］. Ueguchi-Tanaka et al.［10］results suggest that the interaction of SLR1 with GA-GID1 might close the lid，thereby ensuring that GA will be held in the substrate pocket.The stabilized complex of GA，GID1，and SLR1 may be targeted by GID2，an F-box protein，leading to its degradation by 26S proteasomes through ubiquitination of the SCFGID2 complex.Yasumura et al.［45］show thatthe interaction betweenGID1 and DELLA components from Selaginella kraussiana（a lycophyte）is GA stimulated.In contrast，GID1-like（GLP1）and DELLA components from Physcomitrella patens （a bryophyte）do not interact，suggesting that GA-stimulated GID1-DELLA interactions arose in the land-plant lineage after the bryophyte divergence.Feng et al.［46］show that in the absence of GA，nuclear-localized DELLA proteins accumulate to higher levels，interact with phytochrome-interacting factor 3（PIF3，a bHLH-type transcription factor），In the presence of GA，GID1 proteins （GA receptors）elevate their direct interaction with DELLA proteins in the nucleus.On the contrary，Hirano et al.［47］suggests that the interaction betweenGID1andDELLA proteins seems to be a simple biochemical reaction that does not require additional factors in the nucleus.Yamamoto et al.［48］found that soybean（Glycine max）and Brassica napus also have similar unique GID1s that have GA-independent interaction activity with DELLA，indicating that these GID1s occur in various kinds of dicot species and might have crucial roles in some developmental process.Recent physiologicaland biochemicalstudies of these proteins have enabled us to construct a model of GA signaling:GA perception is mediated by GID1 and GA promotes plant growth by GID1-mediated destabilization of the DELLA proteinvia the 26S proteasome pathway［49-50］.However，PpGID1s do not display GA-binding activity in vitro or interact with PpDELLAs in yeast twohybrid assays［51，20］.In addition，the mutant lacking functions of both DELLALike genes does not show any growth defects［20］.These results suggest that the roles of PpGID1s and PpDELLAs are distinct from those in angiosperms.（Sun，2011）Comparative studies suggestthatthe functionalGA-GID1-DELLA module is highly conserved among vascular plants，but not in the bryophytes.Hauvermale et al.［52］researches show that In proteolytic GA signaling，the SLEEPY1（SLY1）F-box protein targets DELLA proteins in the GID1-GA-DELLA complex for destruction through the ubiquitin-proteasome pathway.Non-proteolytic GA signaling in sly1 mutants where GA cannot target DELLA proteins for destruction，requires GA and GID1 gene function. Based on comparison of gid1 multiple mutants to sly1 gid1 mutants，GID1a is the primary GA receptor stimulating stem elongation in proteolytic and nonproteolytic signaling，and stimulating fertility in proteolytic GA signaling. GID1b plays the primary role in fertility，and a secondary role in elongation during non-proteolytic GA signaling. The stronger role of GID1b in nonproteolytic GA signaling may result from the fact that GID1b has higher affinity for DELLA protein than GID1a and GID1c.

Cloning and Expression of GA Receptors

Although the biosynthesis of GA has been well characterized［4，53］，little is known about how plants perceive GA and how the GA signal is transmitted to cause GA-regulated plant growth.In order to further understand the signal transduction of gibberellic acid in the plants.In recent years，GID1 protein as an important signal recognition and conduction has become another hot topics about signal transduction of GA after the DELLA protein.At present，the GID1 gene of many plants has been cloned and expression analyzed.

Recently，a soluble GA receptor was isolated through analysis of a rice（Oryza sativa）GA insensitive dwarf1（gid1）mutant［5］.Soon afterwards，Nakajima et al.［6］cloned three homologousgenesin Arabidopsis，called GID1a，GID1b，and GID1c，and confirmed that all three genes encode proteins that bind GA in vitro and rescue the GA-insensitive dwarf phenotype when expressed （individually）in the rice gid1 mutant.Sui et al.［54］According to other plants conserved domains of GA acceptor，pairsofdegenerate primers were designed.A fragment with a size of 7 00 base pairs was obtained in maize，and their amino acid had higher similarity with 80.5%in rice and maize.According to several different crops EST sequences，amino acid sequences coding these genes were deduced in sorghum，wheat，maize and cotton［55］.Their amino acid sequences had higher similarity to rice with 81.44%，81.36%，80.50%，79.14%，63.13%，respectively.Dong et al.［56］cloned six cotton GID1 homologous genes（GhGID1-1-6）by searching EST sequences.The research cloned the gibberellin receptor gene DID1 fragment in sugarcane by differential expression［57-58］.Li et al.［59］believe that partial cDNA sequnences encoding gibberellins receptor GID1 was isolated from the normal and mutant catkin of the Castanea mollissima by reverse transcription polymerse chain reaction（RT-PCR），the GID1 partial cDNA is 786 bp，putative amino acids have 91%similarity with GID1 in Populus trichocarpa and 85%in Ricinus communis and Gossypium hirsutum.Liu et al.［60］show that a GA receptor gene，named BnGID1B（GenBank accession HQ589349）was cloned by the method of homologous cloning from Brassica napusL，the cDNA length sequence of its open reading frame （ORF）was 1077 bp，encoding a protein of 358 amino acid residues with 40 203.4 Da molecular weight and a theoretical isoelectric point of 6.26，its genomic DNA sequence contains an intron and two exons.Sequence comparison with Arabidopsis GID1B showed that，BnGID1B gene had 86.3%similarity of nucleotide sequence and 91.64%similarity of the amino acid sequence. Voegele et al.［61］show that three L. sativum GID1 cDNAs were cloned and analysed，including the complete coding sequences（cds）and the 5'and 3' untranslated regions （UTRs），Comparative sequence analysis between the cds of GID1a，GID1b，and GID1c of Arabidopsis and L.sativum showed 91.4%，89.4%，and 91%pairwise identity，which are values similar to known orthologues of L.sativum and Arabidopsis.Ge et al.［62］Basing on sequence comparison of already isolated GA receptors from other plants，7 putative GA receptorGID1genes were identified from the grape genome. Zhang et al.［63］researches show that there probably is only one GID1 with highly conserved sequence in conifers. Although the identity of GID1s is low between conifers and angiosperms，they have highly similar amino acids which are essential for binding affinity to GA and quite conserved functional domains which can interact with DELLA protein.Thus，these orthologous GID1 genes may have functions of re-ceptor.Expression analysis showed that the GID1 had stable expressions in different organs in P.abies and in different developmental stages of male and female cones in Pinus tabuliformis，indicating that GID1 may widely participate in developmental process of these tissues，and the transcriptional regulation of GID1 might not be the key regulatory mechanism in GA signaling pathway.

The Function Identification of GA Receptor Gene

Gibberellins （GAs）are endogenous growth regulators in higher plants.GA-deficientmutants have been largely instrumental in identifying the myriad of developmental processes in which GAs participate，and these mutants have laid basis for identifying and characterizing many of the GA biosynthetic enzymes［4］.Recently，many relevant reports are conducted on the function identification of GA receptor gene and these studies have led to a rapid improvement in our understanding the roles in GA signaling.

In Arabidopsis，three GID1-type GA receptors are known:AtGID1a，AtGID1b，and AtGID1c.After binding to its receptor，the GA-GID1 complex interacts with DELLA proteins which are negative regulators of the GA signalling pathway［1］.Griffiths et al.［21］results show that although single mutants developed normally，gid1a gid1c and gid1a gid1b displayed reduced stem height and lower male fertility，respectively，indicating some functional specificity.Tanaka et al.［64］using a proteomic approach to elucidate the function of gid1，the results showed increased tolerance to cold stress and resistance to blast fungus infection. Iuchi et al.［65］results show that all At-GID1s function as GA receptors in Arabidopsis，but have specific role（s）for growth and development.Willige et al.［41］result show that the atgid1a atgid1b atgid1c triple mutant seeds never germinated voluntarily，and after removing their testae with forceps，the embryos grew but were much shorter than the atgid1a atgid1c double mutant.Suzuki et al.［66］Analyses of multiple loss-off unction mutants for GID1s in Arabidopsis show that any single knock-out mutant of GID1s did not show any visible phenotype，which implies that either or both of the remaining GID1（s）function sufficiently.However，only the atgid1a atgid1c double mutant shows a dwarf phenotype.This finding suggests that AtGID1b does not function effectively in stem elongation.Thus，AtGID1b functions poorly in the stem.Therefore，it supposed that AtGID1c probably functions less effectively than AtGID1a or AtGID1b in the floral organ.

On the other hands，Hirano et al.［22］studies revealed that Sm GID1s have different GA binding properties from GID1s in flowering plants.No evidence was found for the functional conservation of these genes in P. patens，indicating that GID1/DELLA-mediated GAsignaling，if present，differs from thatin vascularplants. Voegele et al.［61］analysis of the Arabidopsisknockoutmutantsforthe three GID1 receptors clearly showed that the AtGID1b receptor is not able to compensate for the phenotype of the gid1agid1c double mutant.However partially overlapping roles are proposed for the GID1ac and GID1b pathways and their downstream target genes during seed germination that may be conserved among Brassicaceae oreven among eudicots. These Studies suggests that a GA-triggered negative feedback loop during seed germination exists forGID1a and GID1c in Brassicaceae seeds，while GID1b-type transcripts are not down-regulation targets.

Conclusions

The GA receptor（GID1）was identified by a combination of biochemical and genetic techniques，the structure of GA receptor have been elucidated. Recently，the biochemical and physiological characteristics of the GA receptor，and interactions between the Gibberellin Receptor with DELLA，and Cloning and expression of GA receptors，its function identification in plant has become the research hot spots. The function of Gibberellin Receptor and interactions between the Gibberellin Receptor with important componemts of the pathways Gibberellin will be important for the study of the central role of in modulating plant growth and development.

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Responsible editor：Xiaoxue WANG

Responsible proofreader：Xiaoyan WU

Supported by Natural Science Foundation of China （31360312），Guangxi Natural Science Foundation（2015GXNSFDA39011），National High Technology Research and Development Program （"863"Program）of China （2013AA102604），National Key Technology R&D Program （2012BAD40B04-3），Guangxi Natural Science Foundation（2014GXNSFBA118087），InternationalScientificCooperation Program ofChina（2013DFA31600），Guangxi Special Fund for Bagui Scholars（2013），Guangxi Academy of Agricultural Sciences project （2014YP03，2014YD02，2015YM13，2015YT03）；Guangxi Academy of Agricultural Sciences Project（2014YP03，2014YD02，2015YM13，2015YT03）.

*Corresponding author.E-mail:liyr@gxaas.net

Received:September 3，2015 Accepted:October 16，2015