Difference between revisions of "Os05g0482400"

From RiceWiki
Jump to: navigation, search
Line 76: Line 76:
 
=== Knowledge Extension ===
 
=== Knowledge Extension ===
 
[[File:HELUYUN1.jpg|right|thumb|150px|''The major GA biosynthesis and catabolism pathways in plants(from reference <ref name="ref13" />).'']]
 
[[File:HELUYUN1.jpg|right|thumb|150px|''The major GA biosynthesis and catabolism pathways in plants(from reference <ref name="ref13" />).'']]
 +
 +
Gibberellins (GAs) are a group of diterpenoid compounds, some of which act as growth-promoting hormones controlling such diverse processes as stem elongation, leaf expansion, seed germination, and flowering. The GA biosynthesis pathway has long been studied, and the majority of genes encoding enzymes in each step of the biosynthesis and catabolism pathways have been identified in the model plant species Arabidopsis thaliana and rice (Oryza sativa)<ref name="ref20" /><ref name="ref21" /><ref name="ref22" />. In rice, some mutants with altered GA metabolism or signaling pathways have been studied in detail<ref name="ref23" /><ref name="ref24" /><ref name="ref25" /><ref name="ref26" />. Recently, GIBBERELLIN INSENSITIVE DWARF1 has been identified as a soluble GA receptor in rice<ref name="ref27" />. Such GA-related mutants have been important not only to identify key components in the GA metabolism and signaling pathways but also to confer useful agronomic traits in cereals, some of which contributed to the success of the green revolution<ref name="ref28" /><ref name="ref29" /><ref name="ref30" />.
 +
 +
GAs are biosynthesized from geranylgeranyl diphosphate, a common C20 precursor for diterpenoids. Conversions of geranylgeranyl diphosphate into bioactive GAs, such as GA1 and GA4, involve three classes of enzymes: plastid-localized terpene cyclases, membrane-bound cytochrome P450 monooxygenases (P450s), and soluble 2-oxoglutarate-dependent dioxygenases (2ODDs). In rice vegetative tissues, the early 13-hydroxylation pathway, leading to the formation of bioactive GA1, has been shown to occur predominantly , while GA4, the bioactive non-13-hydroxylated GA (13-H GA), accumulated to a high level in anthers<ref name="ref31" />.
 +
 +
Bioactive GA1 and GA4 and their immediate precursors GA20 and GA9, respectively, are deactivated by GA 2-oxidases, another class of 2ODDs<ref name="ref32" />. Recently, a new class of 2ODD, including AtGA2ox7 and AtGA2ox8 of Arabidopsis, has been shown to catalyze 2-oxidation of C20-GAs (precursor GAs), including GA12 and GA53<ref name="ref33" /><ref name="ref34" />. In pea (Pisum sativum), a loss-of-function mutation in the PsGA2ox1 gene slender causes a tall phenotype<ref name="ref35" />. However, no loss-of-function mutant of the GA2ox gene family members has been recognized as a tall mutant in other plant species, including Arabidopsis and rice, likely due to functional redundancy among the family members<ref name="ref20" /><ref name="ref26" />. Thus, it has yet to be clarified genetically whether GA 2-oxidation, the only well-characterized GA catabolism reaction so far, is commonly the major GA deactivation step(s) in plants or if any other catabolic route also plays a role in controlling bioactive GA levels.
  
 
==Labs working on this gene==
 
==Labs working on this gene==
Line 117: Line 123:
 
<ref name="ref18">Dong-Lei Yang;Qun Li;Yi-Wen Deng;Yong-Gen Lou;Mu-Yang Wang;Guo-Xing Zhou;Ying-Ying Zhang;Zu-Hua He.(2008) Altered Disease Development in the eui Mutants and Eui Overexpressors Indicates that Gibberellins Negatively Regulate Rice Basal Disease Resistance.Molecular Plant 1(3):528-537.</ref>
 
<ref name="ref18">Dong-Lei Yang;Qun Li;Yi-Wen Deng;Yong-Gen Lou;Mu-Yang Wang;Guo-Xing Zhou;Ying-Ying Zhang;Zu-Hua He.(2008) Altered Disease Development in the eui Mutants and Eui Overexpressors Indicates that Gibberellins Negatively Regulate Rice Basal Disease Resistance.Molecular Plant 1(3):528-537.</ref>
 
<ref name="ref19">Qi, M., and Yang, Y. (2002). Quantification of Magnaporthe grisea during infection of rice plants using real-time polymerase chain reaction and northern blot/phosphoimaging analyses. Phytopathology 92, 870–876.</ref>
 
<ref name="ref19">Qi, M., and Yang, Y. (2002). Quantification of Magnaporthe grisea during infection of rice plants using real-time polymerase chain reaction and northern blot/phosphoimaging analyses. Phytopathology 92, 870–876.</ref>
−
<ref name="ref"> .</ref>
+
<ref name="ref20">Hedden, P., and Phillips, A.L. (2000). Gibberellin metabolism: New insights revealed by the genes. Trends Plant Sci. 5, 523–530.</ref>
−
<ref name="ref"> .</ref>
+
<ref name="ref21">Olszewski,N., Sun,T.-P., and Gubler, F.(2002).Gibberellin signaling:Biosynthesis, catabolism, and response pathways. Plant Cell 14(suppl.), S61–S80.</ref>
−
<ref name="ref"> .</ref>
+
<ref name="ref22">Sun, T.-P., and Gubler, F. (2004). Molecular mechanism of gibberellin signaling in plants. Annu. Rev. Plant Biol. 55, 197–223.</ref>
−
<ref name="ref6"> .</ref>
+
<ref name="ref23">Ueguchi-Tanaka, M., Fujisawa, Y., Kobayashi, M., Ashikari, M.,Iwasaki, Y., Kitano, H., and Matsuoka, M. (2000). Rice dwarf mutant d1, which is defective in the alpha subunit of the heterotrimeric G protein, affects gibberellin signal transduction. Proc. Natl. Acad. Sci.USA 97, 11638–11643.</ref>
−
<ref name="ref6"> .</ref>
+
<ref name="ref24">Ikeda, A., Ueguchi-Tanaka, M., Sonoda, Y., Kitano, H., Koshioka, M.,Futsuhara, Y., Matsuoka, M., and Yamaguchi, J. (2001). slender rice, a constitutive gibberellin response mutant, is caused by a null mutation of the SLR1 gene, an ortholog of the height-regulating gene GAI/RGA/RHT/D8. Plant Cell 13, 999–1010.</ref>
−
<ref name="ref6"> .</ref>
+
<ref name="ref25">Sasaki, A., Itoh, H., Gomi, K., Ueguchi-Tanaka, M., Ishiyama,K., Kobayashi, M., Jeong, D.H., An, G., Kitano, H., Ashikari, M.,and Matsuoka, M. (2003). Accumulation of phosphorylated repressor for gibberellin signaling in an F-box mutant. Science 299,1896–1898.</ref>
−
<ref name="ref6"> .</ref>
+
<ref name="ref26">Sakamoto, T., et al. (2004). An overview of gibberellin metabolism enzyme genes and their related mutants in rice. Plant Physiol.134,1642–1653.</ref>
−
<ref name="ref6"> .</ref>
+
<ref name="ref27">Ueguchi-Tanaka, M., Ashikari, M., Nakajima, M., Itoh, H., Katoh, E.,Kobayashi, M., Chow, T.Y., Hsing, Y.I., Kitano, H., Yamaguchi, I.,
−
<ref name="ref6"> .</ref>
+
and Matsuoka, M. (2005). GIBBERELLIN INSENSITIVE DWARF1 encodes a soluble receptor for gibberellin. Nature 437, 693–698.</ref>
−
<ref name="ref6"> .</ref>
+
<ref name="ref28">Peng, J., et al. (1999). ‘Green revolution’ genes encode mutant gibberellin response modulators. Nature 400, 256–261.</ref>
 +
<ref name="ref29">Sasaki, A., Itoh, H., Gomi, K., Ueguchi-Tanaka, M., Ishiyama, K., Kobayashi, M., Jeong, D.H., An, G., Kitano, H., Ashikari, M., and Matsuoka, M. (2003). Accumulation of phosphorylated repressor for gibberellin signaling in an F-box mutant. Science 299,1896–1898.</ref>
 +
<ref name="ref30">Spielmeyer, W., Ellis, M.H., and Chandler, P.M. (2002). Semidwarf(sd-1), ‘‘green revolution’’ rice, contains a defective gibberellin
 +
20-oxidase gene. Proc. Natl. Acad. Sci. USA 99, 9043–9048.</ref>
 +
<ref name="ref31"> Kobayashi, M., Yamaguchi, I., Murofushi, N., Ota, Y., and Takahashi,N. (1988). Fluctuation and localization of endogenous gibberellins in
 +
rice. Agric. Biol. Chem. 52, 1189–1194.</ref>
 +
<ref name="ref32">Thomas, S.G., Phillips, A.L., and Hedden, P. (1999). Molecular cloning and functional expression of gibberellin 2-oxidases, multifunctional
 +
enzymes involved in gibberellin deactivation. Proc. Natl. Acad. Sci.USA 96, 4698–4703.</ref>
 +
<ref name="ref33">Schomburg, F.M., Bizzell, C.M., Lee, D.J., Zeevaart, J.A.D., and Amasino, R.M. (2003). Overexpression of a novel class of gibberellin
 +
2-oxidases decreases gibberellin levels and creates dwarf plants.Plant Cell 15, 151–163.</ref>
 +
<ref name="ref34"> Lee, D.J., and Zeevaart, J.A.D. (2005). Molecular cloning of GA 2-oxidase3 from spinach and its ectopic expression in Nicotiana
 +
sylvestris. Plant Physiol. 138, 243–254.</ref>
 +
<ref name="ref35"> Lester, D.R., Ross, J.J., Smith, J.J., Elliott, R.C., and Reid, J.B.(1999). Gibberellin 2-oxidation and the SLN gene of Pisum sativum.
 +
Plant J. 19, 65–73.</ref>
 +
<ref name="ref36"> .</ref>
 +
<ref name="ref37"> .</ref>
 +
<ref name="ref38"> .</ref>
 +
<ref name="ref39"> .</ref>
  
  

Revision as of 04:40, 12 June 2014

The rice EUI1 (ELONGATED UPPERMOST INTERNODE) gene controls the growth of the first internode at the top of rice, and the recessive mutations of this site cause the abnormal elongation of internodes at the top of rice.

Annotated Information

Function

EUI gene is a recessive gene which was first reported by Rutger and Carnahan [1], which is characterized by the near doubling length of the uppermost internode, excess panicle exsertion and an increased panicle length in comparison with the wild type.The EUI mutant shows notably rapid elongation of the uppermost internode at the heading stage. Because of its prospective application to amend panicle enclosure in the male-sterile lines, this recessive trait, along with male sterility,maintainer and restorer, was referred to as the fourth genetic element of hybrid rice production [1].

The EUI1 gene was mapped on the central part of the long arm of chromosome 5, and located within two overlapping bacterial artificial chromosome (BAC) clones, OSJNBa0018K15 and OSJNBa0095J22, in an interval of 98 kb between markers M0387 and M01 [2].The EUI1 gene encodes a putative cytochrome P450 protein, CYP714D1. It can control the elongation of internodes and determines the plant height and underlies the grain yield .EUI1 plays a negative role in gibberellinmediated regulation of cell elongation in the uppermost internode of rice [3][4].

Rice internodes are important organs that affect plant height, yield, lodging resistance, and panicle exsertion, and also constitute a unique system for the study of gibberellic acid (GA) signaling. A spontaneous recessive tall mutant in rice called eui, which shows notably rapid elongation of the uppermost internode at the heading stage, was picked up in a japonica variety of rice some years ago [1]. The mutant (zeui stock) is characterized by a near-doubling in the length of the uppermost internode, extreme panicle exsertion, and an increase in panicle length, with little or no effect on other internodes or plant characters. This recessive genotype with the elongated uppermost internode is a very useful trait in hybrid rice seed production, and was referred to as the fourth genetic element of hybrid rice, in addition to the traits male sterility, maintainer and restorer [1]. Panicle enclosure, a phenotype which is typical of almost all male-sterile (MS) lines [5] [6] [7], greatly reduces seed production by hybrid rice, necessitating the use of large amounts of exogenous GA to cause panicle exsertion. However, GA application not only increases the cost of seed production, but also greatly increases the rate of seed germination on the panicle, resulting in decreased quality and shortened storage life of hybrid seeds. In contrast, eui MS plants need little or no exogenous GA application for hybrid seed production[6] [8]. Therefore, the eui mutant represents a breakthrough in hybrid rice production[9].Previous genetic and physiological studies have shown that the eui phenotype is due to a recessive mutation in a single gene, which functions similarly in both japonica and indica subspecies and confers highest sensitivity to exogenously applied GA3 at the seedling and heading stages among the genotypes studied, although the level of GAs in eui plants is already higher than that in wild-type plants [9][10][11]. These results strongly suggested that the Eui gene is likely to be involved in GA metabolism or signaling.

The eui1 mutants have increased cell elongation, higher levels of endogenous active gibberellin, and also a higher sensitivity to exogenously applied GA3
(from reference [3]).
.Expression of genes involved in gibberellin biosynthesis, such as gibberellin-20-oxidase and gibberellin-3-oxidase, was elevated in the uppermost internodes of eui1 mutants in comparison with that of the wild type . In contrast, transgenic rice plants overexpressing EUI1 exhibit typical gibberellin-deficient-like phenotypes, such as dark green leaves and dwarf architecture, which can be partially rescued by exogenous gibberellins applications. Consistent with these observations, overexpression of EUI1 causes increased expression of several gibberellin biosynthesis genes encoding gibberellin-20-oxidase2, gibberellin-20-oxidase3 and gibberellin-3-oxidase2 (D18 gene), that are responsible for the conversion of active gibberellin, but decreased expression of the genes encoding gibberellin-2-oxidase1 and gibberellin-2-oxidase3, that are involved in gibberellin deactivation[3]. In the EUI1-RNAi transgenic plants, the reduction of EUI1 expression led to increased lengths of panicle exsertion and uppermost internode elongation, but not comparable with that in EUI1-null mutants
(from reference [3]).
. The increased lengths of panicle exsertion and uppermost internode elongation in five allelic eui1 mutants varied significantly
(from reference [3]).
. Taking these data together, EUI1 plays a negative role in controlling the elongation of rice internodes during heading, and this negative regulation should be correlated with the EUI1 level or activity[3]. The gibberellin-deficient-like phenotypes in the EUI1 overexpressors can be restored more or less by exogenous gibberellin application, and that the SLR1 protein in the EUI1 overexpressors is accumulated to a high level in comparison with the wild type. It has been shown that the SLR1 protein is triggered to undergo rapid degradation by GA3 treatment[12]. After application of exogenous gibberellin, the SLR1 protein can still be detected at a high level, even after 48 h or 2 weeks treatment, which strongly suggested that EUI1 is a fine tuner for modulating gibberellin biosynthesis and/or modifying the stability of SLR1
(from reference [3]).
. Taking all the data together, it is reasonable to postulate that EUI1, the putative P450 monooxygenase, may work as an enzyme for gibberellins inactivation in gibberellin metabolism[3].
(from reference [13]).

During the seedling and tillering stages, eui plants were morphologically similar to wild-type plants[8].However, at the heading stage, the eui mutant exhibited an extremely elongated uppermost internode, with slightly elongated second and third internodes and panicle. Because of the enhanced internode elongation, the stem exposed between the ear and the flag leaf sheath (panicle exsertion) is much longer in the eui mutant than in wild-type plants . The enhanced internode elongation of the eui mutant was due to longitudinally increased cell lengths but not to an increase in the number of cells (Figure . These observations suggested that the uppermost internode of the eui mutant might accumulate an excessive amount of biologically active GAs or exhibit an enhanced GA sensitivity[13].

(from reference [13]).

EUI catalyzed 16a,17-epoxidation of non-13-hydroxylated GAs. Consistent with the tall and dwarfed phenotypes of the eui mutant and Eui-overexpressing transgenic plants, respectively, 16a,17-epoxidation reduced the biological activity of GA4 in rice, demonstrating that EUI functions as a GA deactivating enzyme[13].

Overexpression of EUI1 gave rise to the gibberellin-deficient-like phenotypes, which could be partially reversed by supplementation with gibberellin .Furthermore, apart from the alteration of expression levels of the gibberellin biosynthesis genes, accumulation of SLR1 protein was found in the overexpressing transgenic plants,indicating that the expression level of EUI1 is implicated in both gibberellin-mediated SLR1 destruction and a feedback regulation in gibberellin biosynthesis.The rice EUI gene encodes a cytochrome P450 monooxygenase that deactivates bioactive gibberellins (GAs)[3].

Eui regulates root starch granule development and gravity responses (from reference [14]).

An early study showed that GA plus kinetin treatment could remove starch granules in amyloplasts and therefore change gravisensitivity of cress roots[15]. Starch granules were almost completely absent in eui root-tip cells, while their generation was enhanced in the roots of Eui-OX plants compared with WT plants. As a consequence of the altered starch granule development, Eui-OX roots were more hypersensitive than the WT to gravity. After 2h of rotation away from vertical, most Eui-OX root tips bended near vertical. All WT and Eui-OX root tips bended vertically when roots were rotated over 12h. These results indicate that Eui is also involved in GA homeostasis in rice roots and reveal a novel role for GA in gravity responses[14].

RNAi of Eui effectively increases internode elongation (from reference [14]).

The eui phenotype that has increased panicle exertion (heading performance) has been used in breeding for male sterile varieties of hybrid rice[6][7][8].Using a double-stranded RNA (dsRNA) transgenic approach[16], more than 80% of independent Eui knockout/knockdown transgenic plants showed an elongated internode phenotype with decreased or undetectable expression of Eui that is similar to the eui mutant. These RNAi lines were stably inherited within generations .Consequently, this study provides a feasible approach to rapidly develop elite eui rice lines, which requires a long breeding term when using conventional breeding practices[14].

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 [14].

Transgenic RNA interference of the Eui gene effectively increased plant height and improved heading performance. Furthermore, 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 Eui in root starch granule development and gravity responses[14].

The mutants displayed significantly longer lesion to PXO99A than their controls. Similar results were obtained with infection by DY89031. Similarly, the representative transgenic lines S73 and S74,in which Eui expression was knocked down with RNAi[14], phenocopied the susceptibility of the eui mutants to both the strains. Slightly enhanced resistance to Xoo in semi-dwarf rice containing the mutant "Green Revolution" gene sd1, in comparison with the near-isogenic tall rice carrying the wild-type Sd1 gene that encodes a GA20 oxidase[17]. Therefore, the loss of function of Eui decreased disease resistance, suggesting that EUI might be a positive modulator of basal disease resistance in rice[18].

Overexpression of Eui increases disease resistance to bacterial blight(from reference [18]).

These dwarf Eui-OX lines exhibited significantly increased disease resistance as compared with the wild-type and the separated negative transgenic plants. Dwarf severity was somehow correlated with resistance degree, since lines OX-21, OX-22, OX-39, and OX-47 were more dwarf and appeared more resistant than OX-7, OX-11, and OX-15. In consistency with disease symptom, a three- to five-fold reduction in bacterial growth was measured in OX-39 compared with the wild-type at 8–14 dpi. These results further support the notion that EUI positively regulates disease resistance against bacterial blight[18].

Overexpression of Eui enhances resistance to rice blast Fungus(from reference [18]).

The RNAi lines S73 and S74 exhibited slightly severer symptoms than the wild-type. For grade 5 disease index(the most susceptible symptom), S73 and S74 displayed 20 and 19%, respectively, in comparison with 9% in the wild-type. Similar reduction of blast resistance was also observed in the eui-1 mutant. In contrast, the overexpression lines OX-7, OX-11, OX-21, and OX-39 exhibited significantly enhanced resistance to the fungus, with 70–83% of grade 0 and 1,in comparison with 18% in the wild-type, while the percentages for grades 4 and 5 were greatly decreased in these lines. The further determination of fungal growth in the host using Southern hybridization with the fungal 28S rDNA probe[19] confirmed that fungal growth was greatly limited in the Eui overexpressors, and slightly increased in the RNAi lines compared with the wild-type. These data demonstrated that Eui also positively regulates resistance to rice blast[18].

Knockout or overexpression of the Elongated uppermost internode (Eui) gene encoding a GA deactivating enzyme compromises or increases, respectively, disease resistance to bacterial blight (Xanthomonas oryzae pv. oyrzae) and rice blast (Magnaporthe oryzae)[18].

Expression

Luo et al.[3] found that EUI1 is expressed in most tested tissues and organs, including the young leaves, young roots, shoot apical meristem, the first and the second elongating internodes, flag leaf and young panicles. EUI1 is expressed at a relatively low level in the first elongating internodes, and the highest expression level was found in the young panicles during heading.

(from reference [3]).

The EUI1 gene is constitutively expressed at a low level in all tested organs, except that it is preferentially expressed in the young panicles during heading. In eui1 mutant plants, the most excessive elongation occurred in the first (uppermost) and second internodes, and therefore it was unexpected that the expression of the EUI1 gene in the first and second elongating internodes was relatively low. It is possible that the substrate of EUI1, which controls internode elongation, could exist mainly in the young panicles and might be transported to the action site, the first and the second internodes[3].

(from reference [13]).

Eui express mainly in elongating or dividing tissues, including the shoot apical meristem, the divisional (intercalary meristem) and elongating zones of internodes, nodes of an elongating stem, and panicle (Figures 9B to 9I and 9L) but not in young seedlings roots, and the flag leaf. Before heading,the Eui express mainly in the divisional and node areas of internodes and young panicles. During the heading stage, the Eui mainly express in the flowering spikelets, the divisional zone,and the node of the uppermost internode[13]).

Evolution

The EUI1 gene encodes a putative cytochrome P450 protein. Cytochrome P450 proteins have been known as hemebinding enzymes with mono-oxygenase activities. Some members of the plant P450 family are known to be involved in biosynthesis of plant growth regulators such as gibberellins,jasmonic acid, auxin and brassinosteroids.The phylogenetic analysis of rice and Arabidopsis cytochrome P450 proteins shows that the EUI1/CYP714D1 protein belongs to the CYP714 subfamily of the CYP72 group. The CYP72 clan seems primarily to have conserved functions involved in plant hormone homeostasis. Three known proteins, CYP735A1, CYP735A2 and CYP72A1, belonging to the CYP72 group exhibit the greatest similarity to the EUI1 protein. CYP735A1 and CYP735A2 have been identified as cytokinin biosynthetic enzymes, and CYP72A1 is a secologanin synthase. According to the sequence similarity and the phylogenetic relationship between EUI1 and the three proteins, the EUI1 gene possibly encodes a putative cytochrome P450 involved in gibberellin homeostasis [3].

Fine mapping

(from reference [4]).

The rice eui1 gene was mapped to two overlapping BAC clones, OSJNBa0095J22 and OSJNBb0099O15, between the markers AC40 and AC46, that were 0.64 cM apart and spanned approximately 152 kb[4].

Clone

(from reference [4]).

The EUI1 gene controlling the upper internode elongation in rice is 9804 bp long, and comprises two exons and one intron. The length of the cDNA is 1931 bp containing a 1734 bp ORF, a 110 bp 5¢-UTR and a 87 bp 3¢-UTR. The ORF encodes an unknown 577 amino acid functional protein,that appears to be a member of the cytochrome P450 family[4].

Knowledge Extension

The major GA biosynthesis and catabolism pathways in plants(from reference [13]).

Gibberellins (GAs) are a group of diterpenoid compounds, some of which act as growth-promoting hormones controlling such diverse processes as stem elongation, leaf expansion, seed germination, and flowering. The GA biosynthesis pathway has long been studied, and the majority of genes encoding enzymes in each step of the biosynthesis and catabolism pathways have been identified in the model plant species Arabidopsis thaliana and rice (Oryza sativa)[20][21][22]. In rice, some mutants with altered GA metabolism or signaling pathways have been studied in detail[23][24][25][26]. Recently, GIBBERELLIN INSENSITIVE DWARF1 has been identified as a soluble GA receptor in rice[27]. Such GA-related mutants have been important not only to identify key components in the GA metabolism and signaling pathways but also to confer useful agronomic traits in cereals, some of which contributed to the success of the green revolution[28][29][30].

GAs are biosynthesized from geranylgeranyl diphosphate, a common C20 precursor for diterpenoids. Conversions of geranylgeranyl diphosphate into bioactive GAs, such as GA1 and GA4, involve three classes of enzymes: plastid-localized terpene cyclases, membrane-bound cytochrome P450 monooxygenases (P450s), and soluble 2-oxoglutarate-dependent dioxygenases (2ODDs). In rice vegetative tissues, the early 13-hydroxylation pathway, leading to the formation of bioactive GA1, has been shown to occur predominantly , while GA4, the bioactive non-13-hydroxylated GA (13-H GA), accumulated to a high level in anthers[31].

Bioactive GA1 and GA4 and their immediate precursors GA20 and GA9, respectively, are deactivated by GA 2-oxidases, another class of 2ODDs[32]. Recently, a new class of 2ODD, including AtGA2ox7 and AtGA2ox8 of Arabidopsis, has been shown to catalyze 2-oxidation of C20-GAs (precursor GAs), including GA12 and GA53[33][34]. In pea (Pisum sativum), a loss-of-function mutation in the PsGA2ox1 gene slender causes a tall phenotype[35]. However, no loss-of-function mutant of the GA2ox gene family members has been recognized as a tall mutant in other plant species, including Arabidopsis and rice, likely due to functional redundancy among the family members[20][26]. Thus, it has yet to be clarified genetically whether GA 2-oxidation, the only well-characterized GA catabolism reaction so far, is commonly the major GA deactivation step(s) in plants or if any other catabolic route also plays a role in controlling bioactive GA levels.

Labs working on this gene

  • College of Life Sciences,Zhejiang University, 310029 Hangzhou, China
  • SHARF and National Laboratory of Plant Molecular Genetics, Shanghai Institute of Plant Physiology and Ecology, Shanghai Institutes for Biological Sciences, Chinese Academy of Sciences, 200032 Shanghai, China
  • College of Life Science, Chinese Northeast Agricultural University, 150030 Harbin, China
  • Chinese National Rice Research Institute,310006 Hangzhou, China
  • College of Life Science,South China Agricultural University,510642 Guangzhou, China
  • National Key Laboratory of Plant Genomics, Institute of Genetics and Developmental Biology, Chinese Academy of Sciences, Beijing 100101, PR China
  • China National Rice Research Institute, Chinese Academy of Agricultural Sciences, Hangzhou 310006, PR China
  • National Key Laboratory of Plant Molecular Genetics, Institute of Plant Physiology and Ecology, Shanghai Institutes for Biological Sciences,Shanghai 200032, PR China
  • College of Life Sciences, Nanjing Agricultural University, Nanjing 210095, PR China
  • The State Key Laboratory of Plant Cell and Chromosome Engineering, Institute of Genetics and Developmental Biology, Chinese Academy of Sciences, Beijing 100101, PR China
  • Institute of Genetics and Crop Breeding,Fujian Agriculture and Forestry University, 350002, Fuzhou, China
  • Hainan Institute of Tropical Agricultural Resources, 572025, Sanya, China
  • RIKEN Plant Science Center, Kanagawa 230-0045, Japan
  • Biotechnology Institute, Zhejiang University, Hangzhou 310029, China
  • Research School of Chemistry, Australian National University, Canberra, ACT 0200, Australia
  • Zhejiang Academy of Agricultural Sciences,Hangzhou 310021, China
  • Institute of Insect Science, College of Agriculture and Biotechnology, Zhejiang University, Hangzhou, China

References

  1. ↑ 1.0 1.1 1.2 1.3 Rutger, J.N. and Carnahan, H.L. (1981) A fourth genetic element to facilitate hybrid cereal production. A recessive tall in rice. Crop Sci. 21: 373–376.
  2. ↑ Xu, Y.H., Zhu, Y.Y., Zhou, H.C., Li, Q., Sun, Z.X., Liu, Y.G., Lin, H.X. and He,Z.H. (2004) Identification of a 98-kb DNA segment containing the rice EUI gene controlling uppermost internode elongation, and construction of a TAC transgene sublibrary. Mol. Gen. Genet. 272: 149–155.
  3. ↑ 3.00 3.01 3.02 3.03 3.04 3.05 3.06 3.07 3.08 3.09 3.10 3.11 3.12 Anding Luo;Qian Qian;Hengfu Yin;Xiaoqiang Liu;Changxi Yin;Ying Lan;Jiuyou Tang;Zuoshun Tang;Shouyun Cao;Xiujie Wang;Kai Xia;Xiangdong Fu;Da Luo;and Chengcai Chu.(2006) EUI1, Encoding a Putative Cytochrome P450 Monooxygenase, Regulates the Internodes Elongation by Modulating GA Responses in Rice.Plant and Cell Physiology 47(2): 181-191.
  4. ↑ 4.0 4.1 4.2 4.3 4.4 Ma Hongli;Zhang Shubiao;Ji Lan;Zhu Hongbo;Yang Shulan;Fang Xuanjun;Yang Rencui.(2006) Fine Mapping and in silico Isolation of the EUI1 Gene Controlling Upper Internode Elongation in Rice.Plant Molecular Biology 60(1): 87-94.
  5. ↑ Shen ZT, Yang CD, He ZH (1987) Studies on eliminating panicle enclosure in WA type MS line of rice Oryza sativa subsp. indica).Chin J Rice Sci 1:95–99.
  6. ↑ 6.0 6.1 6.2 Shen ZT, He ZH (1989) Interaction between eui gene and WAMS cytoplasm of rice and improvement of panicle exsertion of MS line. SABRAO J 6:753–756.
  7. ↑ 7.0 7.1 Yang RC, Zhang SB, Huang RH, Yang SL, Zhang QQ (2002) Breeding technology of eui hybrids of rice. Scientia Agricultura Sinica 35:233–237.
  8. ↑ 8.0 8.1 8.2 He ZH, Shen ZT (1994) Sensitivity of elongated internode gene to GA3 and improvement of MS line in rice. Acta Agronomica Sinica 20:161–167.
  9. ↑ 9.0 9.1 Zhang QQ, Yang RC (2003) The effect of different eui genes on biological characters of e-hybrid rice. Scientia Agricultura Sinica 36:735–739.
  10. ↑ He ZH, ETOH K, Shi CH, Shen ZT (1993) Sensitivity of plant height genes to GA3 and their relationship with enzymes in rice. Chin J Rice Sci 7:143–147.
  11. ↑ He ZH, Shen ZT, Li DB (1994) Relations of plant height genes to the sensitivity of GA3 and to the regulation of endogenous hormones in different rice growth stages. Plant Physiol Commun 30:170–174.
  12. ↑ Itoh, H., Ueguchi-Tanaka, M., Sato, Y., Ashikari, M. and Matsuoka, M. (2002)The gibberellin signaling pathway is regulated by the appearance and disappearance of SLENDER RICE1 in nuclei. Plant Cell 14: 57–70.
  13. ↑ 13.0 13.1 13.2 13.3 13.4 13.5 13.6 Yongyou Zhu;Takahito Nomura;Yonghan Xu;Yingying Zhang;Yu Peng;Bizeng Mao;Atsushi Hanada;Haicheng Zhou;Renxiao Wang;Peijin Li;Xudong Zhu;Lewis N. Mander;Yuji Kamiya;Shinjiro Yamaguchi;Zuhua He.(2006)ELONGATED UPPERMOST INTERNODEEncodes a Cytochrome P450 Monooxygenase That Epoxidizes Gibberellins in a Novel Deactivation Reaction in Rice.The Plant Cell 18(2): 442-456.
  14. ↑ 14.0 14.1 14.2 14.3 14.4 14.5 14.6 Yingying Zhang;Yongyou Zhu;Yu Peng;Dawei Yan;Qun Li;Jianjun Wang;Linyou Wang;Zuhua He.(2008)Gibberellin homeostasis and plant height control by EUI and a role for gibberellin in root gravity responses in rice.Cell Research 18(3): 412-421.
  15. ↑ Busch MB, Sievers A.(1990).Hormone treatment of roots causes not only a reversible loss of starch but also of structural polarity in statocytes. Planta 181:358-364.
  16. ↑ Mei C, Zhou X, Yang Y.（2006）Use of RNA interference to dissect defense-signaling pathways in rice. Methods Mol Biol.354:161-172.
  17. ↑ Sasaki,A.,et al.(2002). Green revolution: a mutant gibberellinsynthesis gene in rice. Nature 416, 701–702.
  18. ↑ 18.0 18.1 18.2 18.3 18.4 18.5 Dong-Lei Yang;Qun Li;Yi-Wen Deng;Yong-Gen Lou;Mu-Yang Wang;Guo-Xing Zhou;Ying-Ying Zhang;Zu-Hua He.(2008) Altered Disease Development in the eui Mutants and Eui Overexpressors Indicates that Gibberellins Negatively Regulate Rice Basal Disease Resistance.Molecular Plant 1(3):528-537.
  19. ↑ Qi, M., and Yang, Y. (2002). Quantification of Magnaporthe grisea during infection of rice plants using real-time polymerase chain reaction and northern blot/phosphoimaging analyses. Phytopathology 92, 870–876.
  20. ↑ 20.0 20.1 Hedden, P., and Phillips, A.L. (2000). Gibberellin metabolism: New insights revealed by the genes. Trends Plant Sci. 5, 523–530.
  21. ↑ Olszewski,N., Sun,T.-P., and Gubler, F.(2002).Gibberellin signaling:Biosynthesis, catabolism, and response pathways. Plant Cell 14(suppl.), S61–S80.
  22. ↑ Sun, T.-P., and Gubler, F. (2004). Molecular mechanism of gibberellin signaling in plants. Annu. Rev. Plant Biol. 55, 197–223.
  23. ↑ Ueguchi-Tanaka, M., Fujisawa, Y., Kobayashi, M., Ashikari, M.,Iwasaki, Y., Kitano, H., and Matsuoka, M. (2000). Rice dwarf mutant d1, which is defective in the alpha subunit of the heterotrimeric G protein, affects gibberellin signal transduction. Proc. Natl. Acad. Sci.USA 97, 11638–11643.
  24. ↑ Ikeda, A., Ueguchi-Tanaka, M., Sonoda, Y., Kitano, H., Koshioka, M.,Futsuhara, Y., Matsuoka, M., and Yamaguchi, J. (2001). slender rice, a constitutive gibberellin response mutant, is caused by a null mutation of the SLR1 gene, an ortholog of the height-regulating gene GAI/RGA/RHT/D8. Plant Cell 13, 999–1010.
  25. ↑ Sasaki, A., Itoh, H., Gomi, K., Ueguchi-Tanaka, M., Ishiyama,K., Kobayashi, M., Jeong, D.H., An, G., Kitano, H., Ashikari, M.,and Matsuoka, M. (2003). Accumulation of phosphorylated repressor for gibberellin signaling in an F-box mutant. Science 299,1896–1898.
  26. ↑ 26.0 26.1 Sakamoto, T., et al. (2004). An overview of gibberellin metabolism enzyme genes and their related mutants in rice. Plant Physiol.134,1642–1653.
  27. ↑ Ueguchi-Tanaka, M., Ashikari, M., Nakajima, M., Itoh, H., Katoh, E.,Kobayashi, M., Chow, T.Y., Hsing, Y.I., Kitano, H., Yamaguchi, I., and Matsuoka, M. (2005). GIBBERELLIN INSENSITIVE DWARF1 encodes a soluble receptor for gibberellin. Nature 437, 693–698.
  28. ↑ Peng, J., et al. (1999). ‘Green revolution’ genes encode mutant gibberellin response modulators. Nature 400, 256–261.
  29. ↑ Sasaki, A., Itoh, H., Gomi, K., Ueguchi-Tanaka, M., Ishiyama, K., Kobayashi, M., Jeong, D.H., An, G., Kitano, H., Ashikari, M., and Matsuoka, M. (2003). Accumulation of phosphorylated repressor for gibberellin signaling in an F-box mutant. Science 299,1896–1898.
  30. ↑ Spielmeyer, W., Ellis, M.H., and Chandler, P.M. (2002). Semidwarf(sd-1), ‘‘green revolution’’ rice, contains a defective gibberellin 20-oxidase gene. Proc. Natl. Acad. Sci. USA 99, 9043–9048.
  31. ↑ Kobayashi, M., Yamaguchi, I., Murofushi, N., Ota, Y., and Takahashi,N. (1988). Fluctuation and localization of endogenous gibberellins in rice. Agric. Biol. Chem. 52, 1189–1194.
  32. ↑ Thomas, S.G., Phillips, A.L., and Hedden, P. (1999). Molecular cloning and functional expression of gibberellin 2-oxidases, multifunctional enzymes involved in gibberellin deactivation. Proc. Natl. Acad. Sci.USA 96, 4698–4703.
  33. ↑ Schomburg, F.M., Bizzell, C.M., Lee, D.J., Zeevaart, J.A.D., and Amasino, R.M. (2003). Overexpression of a novel class of gibberellin 2-oxidases decreases gibberellin levels and creates dwarf plants.Plant Cell 15, 151–163.
  34. ↑ Lee, D.J., and Zeevaart, J.A.D. (2005). Molecular cloning of GA 2-oxidase3 from spinach and its ectopic expression in Nicotiana sylvestris. Plant Physiol. 138, 243–254.
  35. ↑ Lester, D.R., Ross, J.J., Smith, J.J., Elliott, R.C., and Reid, J.B.(1999). Gibberellin 2-oxidation and the SLN gene of Pisum sativum. Plant J. 19, 65–73.

Cite error: <ref> tag with name "ref36" defined in <references> is not used in prior text.
Cite error: <ref> tag with name "ref37" defined in <references> is not used in prior text.
Cite error: <ref> tag with name "ref38" defined in <references> is not used in prior text.
Cite error: <ref> tag with name "ref39" defined in <references> is not used in prior text.

Structured Information

Gene Name

Os05g0482400

Description

Cytochrome P450 family protein

Version

NM_001062401.1 GI:115464532 GeneID:4339131

Length

9805 bp

Definition

Oryza sativa Japonica Group Os05g0482400, complete gene.

Source

Oryza sativa Japonica Group 

 ORGANISM  Oryza sativa Japonica Group
           Eukaryota; Viridiplantae; Streptophyta; Embryophyta; Tracheophyta;
           Spermatophyta; Magnoliophyta; Liliopsida; Poales; Poaceae; BEP
           clade; Ehrhartoideae; Oryzeae; Oryza.
Chromosome

Chromosome 5

Location

Chromosome 5:23775727..23785531

Sequence Coding Region

23775837..23776191,23784066..23785444

Expression

GEO Profiles:Os05g0482400

Genome Context

<gbrowseImage1> name=NC_008398:23775727..23785531 source=RiceChromosome05 preset=GeneLocation </gbrowseImage1>

Gene Structure

<gbrowseImage2> name=NC_008398:23775727..23785531 source=RiceChromosome05 preset=GeneLocation </gbrowseImage2>

Coding Sequence

<cdnaseq>atggagagcttcttcgtcttcttcacggcggcggcgttgccggtggtggtggcggcggcggtgatcgccgggctgtgcattacggcggcgtggctggcgaggccgcggcgcgtggcggaggtgttccggaggcaggggatcgacggcccgccgccgtcgtcgttcctggcggggaacctcccggagatgaaggcgagggtggccgccgcggcgtcggcggcggcgccaacggcggacggggaggagaccgcctccgccggcggcggcggcggtggccgggacttcgagaaggacgggttcgacgactactgcaccaggatcttcccttacttccacaagtggaggaaagcctacggcgagacgtacctgtactggctgcggcggcggccggcgctgtacgtgacggacccggagctcatcggcgagatcgggcggtgcgtgtcgctcgacatgggcaagcccaagtacctccagaaaggccaggagccactcttcggcggcggcgtcctcaaggccaacggcgcgtgctgggcgcgccagcgcaaggtcatcgcgccggagttctacatggcccgtgtcagggccatggtccagctcatggtcgacgccgcgcagccgctgatcgcctcctgggaatccaggatcgacgccgctggaggcgcggcggcggcggaggtcgtcgtcgacggcgacctccggagcttctccttcgatgtgatatcgcgggcttgctttgggagtgattactcgagggggagggagatcttcctccgcctccgtgagctgtccgggctcatgtcggagaccagcgtcatcttcagcatcccttcgctgaggcacctgccgacggggaagaaccggaggatctggaggctcacgggggagatccggtcgctgatcatggagctcgtcagggagcggaggtgcgcggcgagggcggcgagggagcacggcgggaaggcggcgccgccgtcgccgccggagcgcgacttcctcggctccatcatcgagaacagcggcgggcagccgcggccggacgacttcgtggtggacaactgcaagaacatctacttcgccgggcacgagacgagcgcggtcaccgcgacgtggtgcctcatgctgctcgccgcgcacccggagtggcaggaccgcgcccgcgccgaggtgctcgaggtctgcggcggcgacggcgccgccgcccccgccgcgccggacttcgacatggtgtcccggatgcggacggtggggatggtggtgcaggaaacgctgcggctgttcccgccgtcgtcgttcgtggtgcgggagacgttccgggacatgcagttgggtaggctgctggcgcccaagggcacctacctgttcgtcccggtgtccaccatgcaccacgacgtcgccgcctggggcccgacggcgaggctgttcgacccgtcccgcttccgcgacggcgtggcggcggcgtgcaagcacccgcaggcgtcgttcatgccgttcggcctcggcgcccgcacctgcctcggccagaacctcgcgctcgtcgaggtcaagacgctcgtcgccgtcgtcctcgcccggttcgagttcacgctctcgccggagtacaggcactcgccggcgttccggctcatcatcgagccggagttcggcctccgcctccgcatccgccgcgccggcggtcaggacgccacgtcacaagttgacacatctactgcacccgtgcatagttctcataattaa</cdnaseq>

Protein Sequence

<aaseq>MESFFVFFTAAALPVVVAAAVIAGLCITAAWLARPRRVAEVFRR QGIDGPPPSSFLAGNLPEMKARVAAAASAAAPTADGEETASAGGGGGGRDFEKDGFDD YCTRIFPYFHKWRKAYGETYLYWLRRRPALYVTDPELIGEIGRCVSLDMGKPKYLQKG QEPLFGGGVLKANGACWARQRKVIAPEFYMARVRAMVQLMVDAAQPLIASWESRIDAA GGAAAAEVVVDGDLRSFSFDVISRACFGSDYSRGREIFLRLRELSGLMSETSVIFSIP SLRHLPTGKNRRIWRLTGEIRSLIMELVRERRCAARAAREHGGKAAPPSPPERDFLGS IIENSGGQPRPDDFVVDNCKNIYFAGHETSAVTATWCLMLLAAHPEWQDRARAEVLEV CGGDGAAAPAAPDFDMVSRMRTVGMVVQETLRLFPPSSFVVRETFRDMQLGRLLAPKG TYLFVPVSTMHHDVAAWGPTARLFDPSRFRDGVAAACKHPQASFMPFGLGARTCLGQN LALVEVKTLVAVVLARFEFTLSPEYRHSPAFRLIIEPEFGLRLRIRRAGGQDATSQVD TSTAPVHSSHN</aaseq>

Gene Sequence

<dnaseqindica>111..465#8340..9718#aagggcgaggtgagcgaggcgagactcgagagagaggagaagatccagaaatcgcaagctgcacgctatagctaggtagctatctccggggaaagagagagtaggctgccatggagagcttcttcgtcttcttcacggcggcggcgttgccggtggtggtggcggcggcggtgatcgccgggctgtgcattacggcggcgtggctggcgaggccgcggcgcgtggcggaggtgttccggaggcaggggatcgacggcccgccgccgtcgtcgttcctggcggggaacctcccggagatgaaggcgagggtggccgccgcggcgtcggcggcggcgccaacggcggacggggaggagaccgcctccgccggcggcggcggcggtggccgggacttcgagaaggacgggttcgacgactactgcaccaggatcttcccttacttccacaagtggaggaaagcctacggtacgtgtacacaagcctctctcctctctactgctcgatttcacatcactagctacatgcatgatgcttgcttgctttgcctgttaattactccgctcatgtgtgcttaattagcttaattaattcacactccggttaatcaagtcatgcaaaacccacaattaatcagctatcaattgagatgagtatataccagaatatatatacttacagtaaatttaataaaaaaataaaaaaaagaaaagaaaagctctctcgatctctctgtgacgatgatacaagaaatttgtggttgccctagtaacaagttgactagagtataaaagctgctaaaatgtcaagagctctccatataaaaaagaacagtgggggagaataattaatactgtcccaaacagcgagaggaagagagggggcaaacgagaaaagagagaataaacaaaattaatgcaggatggaaaccgtggttgggacgaactactcggtttgcttccagtcaaccgtcctgcatcattaagctatggattaataattaattaactgcagtatataaatatatatactcccccacatatattaatcggtcgttggtccaattagacaacagcaaagccatgcctaggtacctgtctaggtaggctatgactgtagtacactcgtactactcacttaatagcacattagtgctcttattttgcgaattaatgaaactttaatcaataataactatatataactataatatatttagtataatttaacaatatatctattcagcgtgctatcgtttcttcagcctatcaatcaacttaaaaaacttagtgttaaattgattttggtgttttcttatcgtaatttttcttcagttttggattgttttttaagttgctaacaattttagttaacaaagctatttggaaggagaggaataatgcatccatgacaaaagtatatgcttcataataattttcatataccagatcttgcccctggtggtgacgtgttaccaaataccgactggaattaatttcttctctctttctttttctctctgccggtgcaatgaggcgacatttttgagttataactgtgcgtatgaatgcatgtattaattgctgtttgctgaggttaataggctagcaataccgttgttttgggggaccccgcagtttgacctagtagagtttatcatggacgcagactgcccctacctgttcttcttcacggacagcatgattaattagctatactagtggtagtactcgatgtcattaataattggtctctccgtacgtgaataattatcgatttttgactttgacaagtactccctccgtacttgttaaggaagtcgttttagacagcaacacggtctccaaaacacaactttaacttcttatttctataaaaatatttattgaaaagtgatatatgtatacttttatgaaatattttttaagacaaatatattcatataatttttacattttaaaacttaatggaaagttacttttccaagatttgactcaaacattgttctaaacgatttcctttatgagtacggaaaaagtatgttttttcactgttcttatgaagtgtacatggaaagtgatcgaatagctctgagcagtactctaggtagggaattacttttagttgcgtacaaatctaacaattaattaatatgccatttttagtgaacataattgtatattttctccgtttcacaatgtaagacattctagcattttccacatgcatattaatgttaatatgtctagattcattaacatcaatataaatataagaaatgctagaatgacttacattataaaacggaggaagtaatttatgctagaatgacttacattataaaacggaggaagtaatttagtacatcaccttcgtcgattattacgaaaacatccatcatatcgatcgatatctgtgtatttgctactccagaaaaatattttctgagaccgacccacagatgcatatatatgcacattattgtggcggttttcctttctttttagatcaatcggtggcctgacggccggcccagcttcgattgaaattaagctgaataattattgtggccgtcatgtgctaatcacccatataattaggcgtggttagaacctgactaagctatttaagccggtcagttaggctttcccaaaacggccatcaggccatgcatgttgccattagctgtccgagcacgccggccaaagctcaaccctcgtgatcacatcaactcgatctccccgatgcttatccctttttgcaagcaaccggtatgtggtgcttagataattaatgcaagaattaacagtgtacctcctatcaatttgccgttagctatgcatgtaatggattagaaattctatatgcatgttttcgtctaacagagtttgatcagcagcagtacttagctaactaattattctgtaatttatataactactataagctagagtttaagcagagtatgtaagtatgatattttaactaatttagttcttgtcatatggcactgtagttgaaaccataaaccgatcaattaataggacaaatattcaacctggttgcctcactgcatgcgttgttttagtttgacaaatgcatgcatgtgtatgatattttatttgcttggcccttaataagtatattatgttgaagtcatgcatcttgacacagaaacttggagcaaaaactatataatatatttcaagaaccaaagggcctgaatactagtacttgcctacatcgatcacatgatgctagtgtcaatcttatttaaggtttttatttcttctttaatgtcaaaatggctattgtttaactatctaaaaacaccaataattagaagactcccactagaccatctcatgatgcattgcacgtaggatcgatatatatggcaacattaattattttacttgaacatctaatcgcgttcgctattattcaacttatgttacatgttctcccctaatttaagcaaaggccggtttgaccaacgggaaacaaataattaaattgatcttccaggcccagtacttatattaattacctagtcattaactacattttctccatatttaatttgaatgcttcgttctgaaatatttaaaggccattaacactaacacggcacacaagaaattaatttaaatatttaatttggctagcatgtatacatatatatgttgatatgcatggatatatatatatatatatatatatatatatatatatatatgtatatatatatatatgtatatatatgtatatatgtatgtatatgtatatgtatatgtatatgtgtgtgtgtgtgtgtgtgtgtgacaatagctaactttagatatgtttttttttcgatgatacatggtcccggcctctgcatcacaaatgcacacatccaacgaatatcgtacaaataataataaaaaaagtgatgggacaaaactcccgactcctacataagacttaggactagtgagctaagggtgtgtttagttcaccaaaattggaaatttgattgaaattagaatgatgtgacggaaaagttgaaagtttgtgtgtgtaggaaagttttgatgtgatggaaaatttggaagtttgaagaaaaagtttagaactaaactcggcctaactgaaaacaaatcaattgatatttgttttcagttatgccgagtttagtttgacaaaattaaacgaaaccatacaatatataattacaaatcaataattaattcctaatgctgcttgatgctactagcttctctcctacacagtgccattgactaggtaattgctcctaaccctagctagctaagcatgcagctcgatcgaatcgatcgagcatgaaagcaacaggtcgatcgatagatggtaagccatgacgatcaatatgatcatatcatgcatgcatgacactgtagagtgtgtactacgtactaccaccactccaccaaccatgcatgcattcctcccccctctctcttgcaccttttaaagctagctaggacgtgtggggacttcaatatatttcttgtgcattgtctctcctacaaggattccacatggagatgtgcatatgcatggggttacaccaattaaaagtgcagcccacactctatggattgaaggtgctagctagatctctgtactctatccgtcccaaaatataagcatttttagttataaatccagatagatagttatccagattcgtagctaaaagttattatattttaggatggaggtaatatagtttagatggtggaaattaagtctagtagctatggggaaaaatggcaaaaataaaattacagtatggggttgtactggaaaattttcatttgaatatatctcttcacaaatatgtaagcctaaatttaatctaaatatatatgggcacaaatgaattaaaaaacatattttctatgaacatatatagcaggaataatatgcattcctcgattttattcccttcaagtgttcatatagtacgttaaaagtttgggttaaaaattttaaataatttctttagttattttgaaatggtatgaatgaaataagagcgtatacccttaagtcataaaagatcatccagattgaataaaaaaacaaagttaagataattaagtagatgaatgtaaatgtgcatatacatatcttgtctttctgttgactagttgtctgttcagaaattatatggcggaaaccctaactgaggtgaaaattcatgtgtaccaagacaaaaaaagaaagttctgaaatcacacaaacaatctcataagtagataagatgatagtgaacaaccatgtaaagatcttttccaaatttaactttcatttgtcagatataaaatgacaaattttgaatcagaaaatattttttttgtttgaaatttaccctcgtttcttgatctgaacagtaccactagatacacctagctacatgtatggtcgattgtgataggctcacacacatgctatagcttgatggttgtcatggtggttgcctagttggcaaaagttcgatgtccagctgtgctactatcatgataattggcagcagcctagctagctagttcaaggggacaatattacactggctttaataatttcctcgtcaccaccttttacctcaaaaacatattcttgaaaaaatgaagaatatgatcatttcaccaaaaatcatggactggccatgattaaccaactcattttggatggttttgttggattcctcaaaccagacacatcattctctgtgtcgagaatggagatgacgtgactagcttttatgactttagaaacatactccctccgtctcaaaaaataaactttaggtttctgtgtccaatgtttgatcgtatatcttatttgaaaaaaatatgaaaaaaattaaaaagacaaatcacgcataaagtattaatcatattttatcatctaacaataattcttttcgtgcttgtgattgtgagtattggaattcttttaatatatgctgaatatactaatgagcataaagtataacagactaatcaagtatgtgagttattcaatccttggcaatttccaatacaaatacagacacactatgtatggaacttttttatatatagtctattaaacaatatatctttgagccgtctgaaaccaaattatatgatatttttttagttaatcataccaagatcattaattgagtaggaatataagacttcaagggggaatcaacaatattaattgtttctgagaacaattattcaaagcaattcataataaaagcatatttattttaggagtgggtacacacactatctagcttgtaccatatataagtcaatgaccacgatatatggctagccacctccctccatcatgaaataaattaacctcatacgaatttagatatagaatatgtccagatttattgattaaatcctagattgatttattttaggacgaagagattacctgcctgcctgcactaaccatctgcgacgccgacgtgtcatccatcggtcgacaaaacgtacgcaggccaacacatgcatacgtacacacaattgcccatttgcaccccgctccccatgtccacgtcagacggcgacacgtacacattaacacatgtacacgtcgttttcacgtacgagctaagcaaatccaacgtacagaaaatagttaatttacccggttgccaactgggacacgtacgccggccacgtcacaggtctacgtggcgcggtgtgggccgggctaggagggttaaaagcccacgcttgtgtgctttggtcaactcgtgcacttattactctgaagagttattagccggttcgatgtgtcgcgattactgttgtaattaagtctgtctctgtggcgatatgtacgttgacgatgattaaggtggtgtttggatctagagacttaactttagtctctgtatttagacactaatttaaagtattaaatatagactaattacaaaactaattacataaataaaaacttatttgcgagataattttttttaagcctaattaatccataattagaaaatatttaatgtagcatcacataggataatcatgggttaattaggctcaatagaatcgtctcgcgaattagtctaagattatggatagtttttattaatagtctacgtttaatatatataattagtgtccaaacatccgataagatagagacttaaaagttttaggtggtgtttagatctagggacttaactttagtccctatatttagacactaatttagagtattaaatatagactacttacaaaactaattacataaataaaagctaattcgcgagataatttttttaagcctaattaatatataattagagaatgtttactgtagcatcacatatgctaatcatggattaattaggctcaatagattcgtctcgcgaattagttcaagattatggatgggttttattaataatctacgtttaatatttataattagtgtccaaacatccgatatgatagggacttaaaagttttaatctcatctaaatatggttttagtctcatctaaacagggtctaatagcagaaagttacgtgagattcgtatactcgtacgtacgtatttgcatcggaattttcaaaatcctagactctttctagtactggtggtacagtttggcaaatctggttttttggatggtttcagcatgtttaatttgcagtgtggaattgttgagatgctagagattttttgcagaacttatgtgaaatttaattaggcgtttcaactgtggctaaacgtgaagatatagcgcgctagagacgtgtgtgttggcagtgttgctttgcaatttgcataggctcgtggtgtttggttcagaaatttatttaaggatccggttgttttttttttcattttccttggtcaatgttgtaaagaatccgtagaatcttggcaagaagattctcccaaacctccaggaaagaggagaagattctccggaatcagtggaagtagtgcagtagctattgactgggatttttgtaagcccggtcaataggttcccctgtacccttggagagaccctgcaagtgcacacagatcacgtatatatccacatgcctatatgtgcatagatcacataggcacgcatcaaaagctttttaatcctagctagcttactttaatctttgattattttcaatagaaatttaatgacgtttaacttgattcatcattcggcgacgtctcttgaacatgttaaaggctgggccgaggcccaccaaaaacggaggcccatgagcaaatttcaaattctgactcgcgcgcggtggcggcaaaattttgcaggcgagacgtacctgtactggctgcggcggcggccggcgctgtacgtgacggacccggagctcatcggcgagatcgggcggtgcgtgtcgctcgacatgggcaagcccaagtacctccagaaaggccaggagccactcttcggcggcggcgtcctcaaggccaacggcgcgtgctgggcgcgccagcgcaaggtcatcgcgccggagttctacatggcccgtgtcagggccatggtccagctcatggtcgacgccgcgcagccgctgatcgcctcctgggaatccaggatcgacgccgctggaggcgcggcggcggcggaggtcgtcgtcgacggcgacctccggagcttctccttcgatgtgatatcgcgggcttgctttgggagtgattactcgagggggagggagatcttcctccgcctccgtgagctgtccgggctcatgtcggagaccagcgtcatcttcagcatcccttcgctgaggcacctgccgacggggaagaaccggaggatctggaggctcacgggggagatccggtcgctgatcatggagctcgtcagggagcggaggtgcgcggcgagggcggcgagggagcacggcgggaaggcggcgccgccgtcgccgccggagcgcgacttcctcggctccatcatcgagaacagcggcgggcagccgcggccggacgacttcgtggtggacaactgcaagaacatctacttcgccgggcacgagacgagcgcggtcaccgcgacgtggtgcctcatgctgctcgccgcgcacccggagtggcaggaccgcgcccgcgccgaggtgctcgaggtctgcggcggcgacggcgccgccgcccccgccgcgccggacttcgacatggtgtcccggatgcggacggtggggatggtggtgcaggaaacgctgcggctgttcccgccgtcgtcgttcgtggtgcgggagacgttccgggacatgcagttgggtaggctgctggcgcccaagggcacctacctgttcgtcccggtgtccaccatgcaccacgacgtcgccgcctggggcccgacggcgaggctgttcgacccgtcccgcttccgcgacggcgtggcggcggcgtgcaagcacccgcaggcgtcgttcatgccgttcggcctcggcgcccgcacctgcctcggccagaacctcgcgctcgtcgaggtcaagacgctcgtcgccgtcgtcctcgcccggttcgagttcacgctctcgccggagtacaggcactcgccggcgttccggctcatcatcgagccggagttcggcctccgcctccgcatccgccgcgccggcggtcaggacgccacgtcacaagttgacacatctactgcacccgtgcatagttctcataattaattggcttctgtaaacagctagtaagtcgtgaactgaaaaaaatacatcacacatacagagacaacagaggattaccattttaccagc</dnaseqindica>

External Link(s)

NCBI Gene:Os05g0482400, RefSeq:Os05g0482400

Retrieved from "https://ngdc.cncb.ac.cn/ricewiki/index.php?title=Os05g0482400&oldid=183979"