Difference between revisions of "Os09g0457900"

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<ref name="ref32">XueW, Xing Y, Weng X, Zhao Y, TangW, Wang L, ZhouH, Yu S, Xu C, Li X,
 
<ref name="ref32">XueW, Xing Y, Weng X, Zhao Y, TangW, Wang L, ZhouH, Yu S, Xu C, Li X,
 
et al (2008) Natural variation in Ghd7 is an important regulator of
 
et al (2008) Natural variation in Ghd7 is an important regulator of
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heading date and yield potential in rice. Nat Genet 40: 761–767
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heading date and yield potential in rice. Nat Genet 40: 761–767.</ref>
 
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Revision as of 05:40, 6 June 2014

The rice gene Os09g0457900,namely OsEATB,is a Ethylene-Response AP2/ERF Factor and restricts ethylene-induced enhancement of gibberellin responsiveness during the internode elongation process by down-regulating the gibberellin biosynthetic gene, ent-kaurene synthase A.[1]

Annotated Information

Function

  • Effect of Transgenic OsEATB on Rice Internode Elongation

Plant height is not only a decisive factor in plant architecture but also an important agronomic trait that is directly linked to yield potential [2][3]. The OsEATB transgenic lines showed dwarf phenotypes, indicating that the internodal elongation process was suppressed by OsEATB overexpression. For experiments, we selected two transgenic lines: 401003 and 401006 (20 individual plants of each line). The transgenic plants and 9311 control plants were cultivated in the same conditions, and plant height was determined at maturity. The average plant heights of the control, 401003, and 401006 lines were 121, 113, and 106 cm, respectively. Thus, 401003 and 401006 plants were 6.61% and 12.40% shorter than the control, respectively (Fig. 3A; Supplemental Table S1). Furthermore, every elongated internode was shortened, especially the fourth internode, which was shortened by an average of 56.30% (Fig. 3A). OsEATB negatively regulated plant height at every growth stage of rice plants. The transgenic plants showed a dwarf phenotype from the three-leaf stage to the mature stage (Fig. 3B; see Fig. 6, A and B, below). The panicles also showed reduced elongation, demonstrating that panicle internodes were shortened as well (see Fig. 6C below). The plant hormone GA is involved in deepwaterresponsive internode elongation [4]. A number of genes encoding GA biosynthetic or signaling pathways have been identified as being associated with this rapid process [3]. To examine the regulation of GA sensitivity in OsEATB transgenic lines, we treated wild-type and transgenic seedlings with 10, 50, and 100 mM GA3 at the four-leaf stage. Under these conditions, both seedlings showed rapid elongation after the 24-h treatments, and OsEATB transgenic seedlings did not show negative sensitivity to GA treatments compared with the control seedlings during this process (Fig. 3, B and C). Together, these results suggested that OsEATB negatively regulates the elongation process of every internode, in every organ, at every growth stage and that the responsiveness of transgenic seedlings to GA is not impaired. Many GAresponsive dwarf plants that are deficient in the biosynthesis of active GAs have been characterized in various plant species [5][6]. These findings, together with our results, led us to conclude that there is negative regulation of GA biosynthesis in the transgenic plants. To test this hypothesis, we used microarray analysis to compare the expression of GA biosynthetic genes between OsEATB transgenic 401006 seedlings and control seedlings at the four-leaf stage. ent-Kaurene is an early intermediate in the GA biosynthesis pathway: ent-kaurene synthase A (CPS) catalyzes the cyclization of geranylgeranyl diphosphate (GGDP) to ent-copalyl diphosphate (CDP), which is then converted to ent-kaurene by ent-kaurene synthase B (KS; Supplemental Fig. S2)[7]. The microarray analysis results showed that the expression of rice CPS OsCPS2 in OsEATB transgenic seedlings was sharply down-regulated (10.32-fold less than the expression level in 9311 wild-type seedlings). In contrast, the expression of GIBBERELLIN 20-OXIDASE (GA20ox2) was 2.77-fold greater in OsEATB transgenic seedlings than in 9311 wild-type seedlings (Table I). GA20ox is one of the major GA biosynthetic genes, and the levels of GAs are homeostatically modulated through negative feedback regulation of GA20ox expression [8]. In addition, no significant differences were found between the OsEATB transgenic seedlings and the wild type in the expression of OsCPS1, OsKS1, OsKO2, and OsKAO. To confirm these results, we used real-time PCR analysis to examine the expression of OsCPS2 and GA20ox2 genes in four-leafstage transgenic line 401006 and 401003 plants and control plants (Fig. 3D). Expression of these two GA biosynthesis-related genes was similarly affected by the overexpression of OsEATB to the microarray results. We also analyzed the expression of OsCPS1, OsKS1, OsKO2, and OsKAO, and in agreement with the microarray data, no significant differences in expression of these genes were found (Fig. 3E).[1]

  • Effect of Rice OsEATB on GA Biosynthesis during Internode Elongation

To examine the negative regulatory relationship between OsEATB and OsCPS2 during internode elongation, we examined the expression of these two genes in elongating internodes of 64- to 73-d-old 9311 plants (Fig. 4A). In elongating internodes of 64-dold plants, OsEATB was expressed at a high level while OsCPS2 was expressed at a much lower level. The expression of OsEATB sharply decreased 3 d later, while that of OsCPS2 increased approximately 10-fold. Expression of OsEATB was remarkably decreased from 67 to 73 d, while expression of OsCPS2 was slightly increased and then remained almost stable. We compared gene expression in elongating internodes of 67-d-old 401006 transgenic and control plants using real-time quantitative PCR analysis. The genes examined were involved in the GA biosynthesis and GA signaling pathways (Fig. 4B). First, we reexamined the expression of OsCPS2 and GA20ox2 at this growth stage. In elongating internodes of transgenic plants, OsCPS2 was down-regulated by 11.21-fold while GA20ox2 was up-regulated by 2.03- fold, compared with their respective expression in wild-type plants. In rice, the expression levels of XET (for xyloglucan endotransglycosylase) and UROD (for uroporphyrinogen decarboxylase) are up-regulated by GA treatment, while that of GOX (for glycolate oxidase) is down-regulated [9]. We investigated the effect of OsEATB on the expression of these three genes. Transgenic lines overexpressing OsEATB showed decreased expression of XET and UROD but slightly increased expression of GOX compared with their respective expression in the wild type. These findings suggested that the GA response was suppressed in the transformants, possibly due to the lack of bioactive GAs. SLENDER RICE1 (SLR1), the only DELLA protein in rice [10] , down-regulates the GA signaling pathway [11] . We evaluated the effect of OsEATB overexpression on this gene and found that expression of SLR1 was down-regulated in transgenic plants. Together, all of these results confirmed that OsEATB negatively regulates the internode elongation process, not via down-regulating GA sensitivity but through restricting GA biosynthesis.[1]

  • Effect of Transgenic OsEATB on Rice Tillering and Panicle Branching

To investigate the function of rice OsEATB on yield traits, we analyzed traits in two transgenic lines (401003 and 401006; 20 individual plants of each line). The transgenic lines and 9311 control plants were cultivated in the same conditions, and we evaluated four yield components: panicles per plant, panicle length, spikelets per panicle, and grain number per panicle. The transgenic plants produced more tillers than the control 9311 lines at the six-leaf stage (Fig. 6B). In rice, the tiller is a specialized grain-bearing branch that forms on the unelongated basal internode. The tiller grows independently of the mother stem (culm) by means of its own adventitious roots [12] . At maturity, transgenic plants had produced 16.95% more panicles than wild-type plants (Fig. 6, A and D; Supplemental Table S1). Spikelets are grass-specific, flower-bearing branches that form on each panicle branch. They consist of primary, secondary, and sometimes higher order panicle branches [13]. We counted the spikelets per panicle in transgenic plants and wildtype lines. The 35S:OsEATB transformants produced more spikelets than control plants (Fig. 6, C and E).We also counted the numbers of primary and secondary branches per panicle. Transgenic lines showed 8.31% more primary spikelets per panicle and 34.41% more secondary branches per panicle compared with the wild type (Fig. 6D; Supplemental Table S1). The finding that 35S:OsEATB transformants producedmore secondary braches, more spikelets per panicle, and more panicles per plant suggested that OsEATB activity is responsible for branching in rice. To investigate the effects of OsEATB on rice grain weight, we examined the 1,000-grain weight at random and observed a slight change. The 1,000-grain weight of the control and transgenic lines is about 30.0 and 25.5g, respectively. Average numbers of grains per plant of the control, 401003, and 401006 lines were 776, 1,096, and 1,254, respectively. The calculated grain weights per plant were about 23.28, 27.95, and 31.98 g, respectively. These results indicated that 401003 and 401006 possess increases in grain yield per plant over the control of 20.06% and 37.37% (Fig. 6E; Supplemental Table S1).[1]

  • OsEATB Is a Novel Rice AP2/ERF

ERFs are an important subfamily of AP2/ERF transcription factors, and they have a variety of functions. Several rice ERF genes have been isolated and identified; for example, OsEBP-89 regulates transcription of the rice Wx gene [14], and OsEREBP1 is related to the regulation of defense responses [15]. In this study, we investigated and confirmed the function of a rice AP2/ERF gene, OsEATB. ERFs can be classified into different subgroups based on their function and group motif. Subgroup I, which includes Arabidopsis (Arabidopsis thaliana) AtERF1 and AtERF2 and tomato (Solanum lycopersicum) Pti4, functions as transcriptional activators. Subgroup II, which includes tobacco (Nicotiana tabacum) NtERF3 and Arabidopsis AtERF3 and AtERF4, functions as transcriptional repressors. However, the functions of ERF subgroups III and IV remain unclear [16]. OsEATB cannot be classified into any of the function-known ERF subgroups (Fig. 1). Our data show that OsEATB encodes a transcriptional factor that localizes to the nucleus and is constitutively expressed in various tissues (Fig. 1). Experimental data from transgenic plants showed that OsEATB plays a crucial role in regulating the rice internode elongation process (Figs. 2–4). Ethylene is perceived by a family of His kinase-like receptors and, downstream, by EIN2, a novel protein containing an integral membrane domain. In the nucleus, the EIN3 family of DNA-binding proteins regulates ethylene-responsive transcription, and an immediate target of EIN3 is the AP2/EREBP family [17]. As a result, most ERFs are regulated by ethylene. Many ERF proteins have been shown to bind to the specific ERE, the GCC box. In our study, we examined the binding activity of OsEATB to the GCC box (Fig. 1). At the same time, we found that the expression level of OsEATB was sharply decreased by ethylene (Fig. 5). Consequently, our results show that this gene is involved in the regulation of ethylene-related. responses. OsEATB overexpression dramatically suppressed the internode elongation process (Fig. 3) and made seedlings hypersensitive to NaCl and ABA. In addition, the expression of this gene was down-regulated by both NaCl and ABA (Fig. 5). Ethylene is mostly reported to trigger the internodal elongation process via GA [11] [18] , and many ERF genes are induced by abiotic stress conditions in rice [19] [16]. Thus, rice OsEATB might be a negative balance regulator of the ethylene-responsive pathway.[1]

  • OsEATB Negatively Regulates Ethylene-Induced

Enhancement of GA Responsiveness by Reducing GA Biosynthesis Plant height is a decisive factor in plant architecture. In rice, there is the potential for rapid internodal elongation, and the degree of elongation determines plant height. This rapid growth response is best demonstrated in deepwater rice, especially in the process of submergence tolerance [20]. Sub1A is an ERF that confers submergence tolerance to rice. This tolerance is mediated by SLR1, which restricts the response to GA. Sub1A augments SLR1 and SLRL1 gene expression, which counteract the elevated responsiveness to GA promoted by the increase in ethylene [21] [11]. The ERF genes SK1 and SK2 allow rice to adapt to deep water. Under deepwater conditions, ethylene accumulates in the plant and induces the expression of these two genes. These SK genes encoding ERFs trigger internode elongation in deepwater rice via GA. In contrast to Sub1A, SK1 and SK2 may stimulate GA responses [18]. Both SK genes and SUB1A encode ERFs and are related toGA, but they have opposing functions in regulating plant height in response to flooding. It is interesting that gene family members in the same subgroup confer different functions related to internode elongation. In conclusion, there is a self-balance of ethylene-induced enhancement of the GA response during the internodal elongation process. It is possible that cross talk mediated by OsEATB between ethylene and GA underlies the differences in rice internode elongation. The OsEATB transgenic seedlings did not show negative sensitivity to GA treatments (Fig. 3), and the expression-level analysis of the genes involved in GA biosynthesis and GA signaling pathways showed that the GA response is suppressed via down-regulating a key enzyme involved in GA biosynthesis and not through up-regulating the important GA sensitivity repressor (Fig. 4). The Arabidopsis GA-deficient mutant dwarf and delayed flowering (ddf1) phenotypes are caused by increased or ectopic expression of a putative AP2 transcription factor, DDF1. DDF1 is involved in the down-regulation of GA biosynthesis [6]. As a potential negative balance regulator of the ethylene-responsive pathway, OsEATB suppresses the internode elongation process through the restriction of GA biosynthesis, specifically downregulating the expression of OsCPS2 (Table I; Fig. 3). The mutation line of OsCPS1 showed a dwarf phenotype without flower or seed development, which is a typical phenotype of GA-deficient rice dwarf mutants [22]. Evidence is presented indicating that OsCPS2 is involved in related secondary metabolism, producing defensive phytochemicals [23].The expression of OsCPS2 is sharply negatively related to OsEATB expression, and the endogenous GA level is decreased in OsEATB transgenic plants alongside suppressed expression of OsCPS2 (Table II). No significant differences between the OsEATB transgenic seedlings and the wild type in the expression of OsCPS1, OsKS1, OsKO2, and OsKAO were found, and the expression of GA20ox2 was slightly up-regulated for the feedback of GA deficiency. All these experimental results strongly demonstrated that OsCPS2 is also associated with GA biosynthesis. Promoter regions (1,300 bp upstream of the translation site) of OsCPS2 do not contain the GCC box motif, suggesting that this gene may not be a direct target of OsEATB. We propose a model for OsEATB-dependent hormonal regulation of internode elongation in rice (Fig. 7). It was suggested that the important function of OsEATB is to negatively regulate the ethylene-induced enhancement of GA responsiveness during the internode elongation process by decreasing GA biosynthesis.[1]

  • OsEATB Regulates Rice Yield Components through the Promotion of Tillering and Panicle Branching

Food security for the ever-increasing world population largely relies on the grain yield of crop plants [24]. The critical components to determine rice yield include grain number and grain weight. Grain number is contingent on the number of spikelets per panicle and the number of panicles per plant. The number of panicles is mainly determined by the plant architecture and the spikelets per panicle by panicle morphology (i.e. the number of primary/secondary/ tertiary branches on each panicle)[25]. Overexpression of OsEATB decreased plant height (Fig. 3) and increased the numbers of panicles per plant and spikelets per panicle (Fig. 6). More panicles and spikelets resulted in a 37.37% increase in grain yield, notwithstanding that the 1,000-grain weight was 15.00% lower in transgenic lines than in the control (Fig. 6). The results of this study show that OsEATB, as a member of the AP2/ERF family, positively regulates rice yield components through the promotion of rice tillering and panicle branching. The basic structure of a rice panicle is determined by the pattern of branch formation. In rice, the MONO CULM1 (MOC1) and LAX PANICLE genes are necessary for branch meristem formation [13]. MOC1 positively regulates tillering by promoting axillary meristem outgrowth [26][27]. Although the molecular mechanisms that underlie the cross talk between plant height and branching are poorly understood, it is well known that rice plant height is strongly negatively correlated with tiller number [28][29]. Higher yields are typically obtained from dwarf crops [30]. The finding that transgenic rice plants harboring the MOC1 gene are dwarf but produce more tillers than wild-type plants [26] provides a good opportunity to investigate the genetic control network. Our results indicate that overexpression of the OsEATB gene (Fig. 2) reduces rice plant height and panicle length at maturity, promoting rice branching potential in both tillers and spikelets, possibly via the regulation of both shoot elongation and axillary outgrowth. The short stature reflects the decreased growth of the mother stem, which allows the growth of more tillers, leaves, and panicles and enhances the energy utilization ratio and biomass production. The functions of OsEATB in regulating rice plant architecture include its effects on plant height (decreased internode elongation) and panicle morphology (increased tiller formation). Studies on the rice AP2/ERF gene OsEATB provide an opportunity to identify agriculturally important functions that can be used to improve rice yield components. In addition, this gene provides a model for investigating cross talk between ethylene and GA in the internodal elongation process.[1]

Expression

  • Expression of OsEATB in Transgenic Rice Lines

To investigate the function of OsEATB, we introduced the plasmid 35S:OsEATB containing the OsEATB gene (Fig. 2A) into indica variety 9311. We produced transgenic plants overexpressing the sense strand of OsEATB. 35S:OsEATB transformants were screened on antibiotic selection medium containing hygromycin. The transgenic plants were checked by PCR using genomic DNA as the template and specific primers (see “Materials and Methods”). Eight independent transformants (T0), regenerated fromhygromycinresistant calli, were shown to contain OsEATB. Eight PCR-positive T1 lines (20 individual plants of each line) shared the same dwarf phenotype. Two transgenic lines (401003 and 401006) were selected for Southern hybridization analysis. A single, specific band of OsEATB transgene was observed in 35S: OsEATB transgenic lines 401003 and 401006 (Fig. 2B). These results showed that OsEATB was integrated into the rice genome. The expression level of the OsEATB gene was examined in three-leaf-stage transgenic plants and control plants by real-time PCR analyses (Fig. 2C). Expression of OsEATB was 9.77-fold greater in the 401006 transgenic plants and 4.00-, 2.83-, and 2.00-fold greater in the 401003, 401005, and 401007 transgenic plants than in 9311 control plants, respectively. These results were further confirmed by RNA hybridization.[1]

  • Overexpression of OsEATB Decreases the Endogenous

GA Level in Rice CPS catalyzes the cyclization of GGDP to CDP, which is then converted to ent-kaurene before going through a series of processes to produce GA and various GA derivatives. Since OsEATB represses the expression of OsCPS2, we determined the endogenous GA level in three-leaf-stage 401006 OsEATB transgenic plants and 9311 control plants, six-leaf-stage 401006 and 401003 OsEATB transgenic plants and 9311 control plants. As shown in Table II, the endogenous levels of all GAs tested (i.e. GA12, GA53, GA24, GA19, GA9, GA20, GA4, and GA1) were significantly lower in OsEATB transgenic plants than in the wild type, while the levels of abscisic acid (ABA) and indole-3-acetic acid (IAA) were not significantly changed. Therefore, GA levels are decreased along with the suppression of OsCPS2 expression (Supplemental Fig. S2).[1]

  • OsEATB Expression Is Negatively Regulated by Ethylene, ABA, and Abiotic Stress in Rice

OsEATB binds to the ERE, the GCC box, resulting in decreases in the levels in GAs. While increased responsiveness to GA is often associated with an increase in ethylene [18], differential expression of OsEATB was observed in 18-d-old (three-leaf stage) rice 9311 seedlings in response to 50 mM ethephon treatment (Fig. 5A). Ethephon treatment sharply decreased the expression of OsEATB, and expression remained at low levels for 24 h after treatment. Because the expression level of OsEATB was down-regulated by ethephon treatment and OsCPS2 is decreased by OsEATB, we examined the effect of ethylene treatment on OsCPS2. Real-time quantitative PCR analysis was used to investigate the expression of OsEATB and OsCPS2 at 0, 3, 6, 12, and 24 h of ethephon treatment (Fig. 5B). Expression of OsCPS2 increased while that of OsEATB was sharply down-regulated by the treatment. Nevertheless, after 12 h of treatment, the expression of OsCPS2 was stabilized to a normal level via the self-balance regulation of other pathways, for it is a key enzyme in GA biosynthesis. This result indicated that an increase in the amount of OsCPS2 in response to ethylene is correlated with the decrease in OsEATB expression responding to ethylene. In rice, most ERF genes are induced by abiotic stress conditions [16]. To investigate the effect of transgenic OsEATB on the response to abiotic stress in rice, 18-d-old seedlings of 9311 control and transgenic 401006 plants were treated with 200 mM NaCl or 100 mM ABA (Fig. 5, C and D). The transgenic seedlings treated with 200 mM NaCl showed wilting and rolling of leaves, while 9311 control plants showed a significantly higher survival rate (Fig. 5C). After 48 h of stress followed by 48 h of watering, almost all of the transgenic plants never recovered, whereas 42% of the control plants survived. The transgenic seedlings showed stronger chlorosis in response to the 100 mM ABA treatment, while the chlorosis of the 9311 control line was slower (Fig. 5D). Thus, overexpression of OsEATB made seedlings hypersensitive to NaCl and ABA. We also analyzed whether the expression of OsEATB was regulated by these two conditions. Seedlings (18 d old) of 9311 subjected to a salt treatment (200 mM NaCl in Hoagland solution) showed rapid down-regulation of OsEATB expression within 24 h after the treatment compared with that in the watertreated control plants (Fig. 5A). Moreover, expression of OsEATB was slightly decreased in 18-d-old 9311 seedlings in response to ABA treatment (100 mM ABA in Hoagland solution) compared with that in the water-treated control plants (Fig. 5A). OsERF3 is upregulated by ethylene treatment[31] , while ABA treatment induces the expression of CPD [32]). Therefore, we chose OsERF3 as a positive control for ethylene responsiveness and CPD as a positive control for ABA responsiveness. These results strongly suggest that OsEATB is responsive to ethylene and environmental stresses and, therefore, may be involved in the regulation of ethylene-related responses.[1]

Evolution

  • There are 161 known AP2/ERFs in indica rice, 103 of

which are potential ERFs with a single complete AP2 domain (http://plntfdb.bio.uni-potsdam.de). Of these 103 potential ERFs, 38 cannot be classified into any of the four functional subgroups [16], for they lack typical group motifs. We selected OsEATB from these 38 ERFs for further investigation because, like the members of subgroup IV, it contains a nuclear localization signal adjacent to the highly conserved AP2 domain (Fig. 1A; Supplemental Fig. S1). Multiple sequence alignment of OsEATB with other known ERF proteins showed that their similarity was restricted to the DNA-binding domain region, and OsEATB could not be classified into any of the ERF subgroups with previously defined functions (Fig. 1B). We inferred that OsEATB is a rice ERF that might have a potential new function. Based on the data from the National Center for Biotechnology Information GenBank database (http:// www.ncbi.nlm.nih.gov/), OsEATB has no intron and exists as a single-copy gene. Using PCR, we isolated and cloned its full-length 825-bp open reading frame (ORF) from genomic DNA of indica variety 9311. Sequence data for this article have been deposited at GenBank under accession number EU622934. The 825-bp ORF encodes a protein consisting of 274 amino acids.We used the yeast one-hybrid system and electrophoretic mobility shift assays to examine the binding activity of OsEATB to the GCC box. As shown in Figure 1C, the first section of each plate shows the yeast reporter strain harboring pHIS-GCC box and pGAD-OsEATB, while the second section shows yeast cells harboring pHIS-muGCC box and pGADOsEATB. The third section of each plate shows yeast cells transformed with pHIS-GCC box and the negative control vector pGAD242, and the fourth section shows yeast cells transformed with pGAD-OsEATB and negative control vector pHIS2.1. The yeast reporter strain containing pHIS-GCC box and pGADOsEATB grew well on both synthetic dextrose (SD)/ Trp2Leu2 medium and SD/Trp2Leu2His2/50 mM 3-amino-1,2,4-triazole (3-AT) medium. The other three yeast reporter strains grew well on SD/Trp2Leu2 medium but not on SD/Trp2Leu2His2/50 mM 3-AT medium. The band corresponding to the GCC boxcontaining labeled probe and purified His fusion recombinant OsEATB protein complex showed a marked mobility shift compared with the free probe band (Fig. 1D). And the muGCC box-containing labeled fragment served as a competitor. These results demonstrated that OsEATB could specifically bind to the GCC box. If OsEATB acts as a transcription factor, then its nuclear localization signal should localize it to the nucleus. To confirm the subcellular localization of OsEATB, we fused the coding sequence of OsEATB with that of GFP under the control of the 35S promoter (35S:OsEATB:GFP) and delivered the construct into onion (Allium cepa) epidermal cells by particle bombardment (Fig. 1E). The OsEATB protein localized the GFP signal to the nucleus (Fig. 1F).We investigated the tissue expression pattern of OsEATB by reverse transcription (RT)-PCR using rice total mRNA from roots, culms, leaves, and young panicles as the template. The results showed that OsEATB mRNA is expressed constitutively in these four tissues and is expressed at higher levels in roots and leaves than in culms and young panicles (Fig. 1G).[1]

Labs working on this gene

  • State Key Laboratory of Genetic Engineering, Institute of Genetics, School of Life Sciences, Fudan University,Shanghai 200433, China.
  • Key Laboratory of Analytical Chemistry for Biology and Medicine, Department of Chemistry, Wuhan University, Wuhan 430072, China.

References

  1. ↑ 1.00 1.01 1.02 1.03 1.04 1.05 1.06 1.07 1.08 1.09 1.10 Weiwei Qi, Fan Sun, Qianjie Wang, Mingluan Chen, Yunqing Huang, Yu-Qi Feng, Xiaojin Luo, and Jinshui Yang(2011) Rice Ethylene-Response AP2/ERF Factor OsEATB Restricts Internode Elongation by Down-Regulating a Gibberellin Biosynthetic Gene. Plant Physiol 157:216-228.
  2. ↑ Huang N, Courtois B,Wang GL (1996) Association of quantitative trait loci for plant height with major dwarfing genes in rice. Heredity 77: 130–137
  3. ↑ 3.0 3.1 Yang XC, Hwa CM (2008) Genetic modification of plant architecture and variety improvement in rice. Heredity 101: 396–404
  4. ↑ Raskin I, Kende H (1984) Role of gibberellin in the growth response of submerged deep water rice. Plant Physiol 76: 947–950
  5. ↑ Ross JJ, Murfet IC, Reid JB (1997) Gibberellin mutants. Physiol Plant 100: 550–560
  6. ↑ 6.0 6.1 Magome H, Yamaguchi S, Hanada A, Kamiya Y, Oda K (2004) dwarf and delayed-flowering 1, a novel Arabidopsismutant deficient in gibberellin biosynthesis because of overexpression of a putative AP2 transcription factor. Plant J 37: 720–729
  7. ↑ Yamaguchi S, Sun T, Kawaide H, Kamiya Y (1998) The GA2 locus of Arabidopsis thaliana encodes ent-kaurene synthase of gibberellin biosynthesis. Plant Physiol 116: 1271–1278
  8. ↑ Xu YL, Li L,Wu K, Peeters AJ, Gage DA, Zeevaart JA (1995) The GA5 locus of Arabidopsis thaliana encodes a multifunctional gibberellin 20-oxidase: molecular cloning and functional expression. Proc Natl Acad Sci USA 92: 6640–6644
  9. ↑ Yang GX, Jan A, Shen SH, Yazaki J, Ishikawa M, Shimatani Z, Kishimoto N, Kikuchi S, Matsumoto H, Komatsu S (2004) Microarray analysis of brassinosteroids- and gibberellin-regulated gene expression in rice seedlings. Mol Genet Genomics 271: 468–478
  10. ↑ Achard P, Genschik P (2009) Releasing the brakes of plant growth: how GAs shutdown DELLA proteins. J Exp Bot 60: 1085–1092
  11. ↑ 11.0 11.1 11.2 Fukao T, Bailey-Serres J (2008) Submergence tolerance conferred by Sub1A is mediated by SLR1 and SLRL1 restriction of gibberellin responses in rice. Proc Natl Acad Sci USA 105: 16814–16819
  12. ↑ Li YH (1979) Morphology and Anatomy of Grass Family Crops. Shanghai Science and Technology Press, Shanghai, China, pp 138–142
  13. ↑ 13.0 13.1 Furutani I, Sukegawa S, Kyozuka J (2006) Genome-wide analysis of spatial and temporal gene expression in rice panicle development. Plant J 46: 503–511
  14. ↑ Zhu Y, Cai XL, Wang ZY, Hong MM (2003) An interaction between a MYC protein and an EREBP protein is involved in transcriptional regulation of the rice Wx gene. J Biol Chem 278: 47803–47811
  15. ↑ Cheong YH, Moon BC, Kim JK, Kim CY, Kim MC, Kim IH, Park CY, Kim JC, Park BO, Koo SC, et al (2003) BWMK1, a rice mitogen-activated protein kinase, locates in the nucleus andmediates pathogenesis-related gene expression by activation of a transcription factor. Plant Physiol 132: 1961–1972
  16. ↑ 16.0 16.1 16.2 16.3 Cao Y, Song F, Goodman RM, Zheng Z (2006) Molecular characterization of four rice genes encoding ethylene-responsive transcriptional factors and their expressions in response to biotic and abiotic stress. J Plant Physiol 163: 1167–1178
  17. ↑ Chang C, Shockey JA (1999) The ethylene-response pathway: signal perception to gene regulation. Curr Opin Plant Biol 2: 352–358
  18. ↑ 18.0 18.1 18.2 Hattori Y, Nagai K, Furukawa S, Song XJ, Kawano R, Sakakibara H,Wu J, Matsumoto T, Yoshimura A, Kitano H, et al (2009) The ethylene response factors SNORKEL1 and SNORKEL2 allow rice to adapt to deep water. Nature 460: 1026–1030
  19. ↑ Thara VK, Tang X, Gu YQ, Martin GB, Zhou JM (1999) Pseudomonas syringae pv tomato induces the expression of tomato EREBP-like genes pti4 and pti5 independent of ethylene, salicylate and jasmonate. Plant J 20: 475–483
  20. ↑ Kende H, van der Knaap E, Cho HT (1998) Deepwater rice: a model plant to study stem elongation. Plant Physiol 118: 1105–1110
  21. ↑ Xu K, Xu X, Fukao T, Canlas P, Maghirang-Rodriguez R, Heuer S, Ismail AM, Bailey-Serres J, Ronald PC, Mackill DJ (2006) Sub1A is an ethylene-response-factor-like gene that confers submergence tolerance to rice. Nature 442: 705–708
  22. ↑ Sakamoto T, Miura K, Itoh H, Tatsumi T, Ueguchi-Tanaka M, Ishiyama K, Kobayashi M, Agrawal GK, Takeda S, Abe K, et al (2004) An overview of gibberellin metabolism enzyme genes and their related mutants in rice. Plant Physiol 134: 1642–1653
  23. ↑ Prisic S, Xu MM, Wilderman PR, Peters RJ (2004) Rice contains two disparate ent-copalyl diphosphate synthases with distinct metabolic functions. Plant Physiol 136: 4228–4236
  24. ↑ XueW, Xing Y, Weng X, Zhao Y, TangW, Wang L, ZhouH, Yu S, Xu C, Li X, et al (2008) Natural variation in Ghd7 is an important regulator of heading date and yield potential in rice. Nat Genet 40: 761–767.
  25. ↑ Zha XJ, Luo XJ, Qian XY, He GM, Yang MF, Li Y, Yang JS (2009) Overexpression of the rice LRK1 gene improves quantitative yield components. Plant Biotechnol J 7: 611–620
  26. ↑ 26.0 26.1 Li XY, Qian Q, Fu ZM, Wang YH, Xiong GS, Zeng DL, Wang XQ, Liu XF, Teng S, Hiroshi F, et al (2003) Control of tillering in rice. Nature 422: 618–621
  27. ↑ Leyser O (2005) The fall and rise of apical dominance. Curr Opin Genet Dev 15: 468–471
  28. ↑ Hong Z, Ueguchi-Tanaka M, Umemura K, Uozu S, Fujioka S, Takatsuto S, Yoshida S, Ashikari M, Kitano H, Matsuoka M (2003) A rice brassinosteroid- deficient mutant, ebisu dwarf (d2), is caused by a loss of function of a new member of cytochrome P450. Plant Cell 15: 2900–2910
  29. ↑ Booker J, Auldridge M, Wills S, McCarty D, Klee H, Leyser O (2004) MAX3/CCD7 is a carotenoid cleavage dioxygenase required for the synthesis of a novel plant signaling molecule. Curr Biol 14: 1232–1238
  30. ↑ Spielmeyer W, Ellis MH, Chandler PM (2002) Semidwarf (sd-1), “green revolution” rice, contains a defective gibberellin 20-oxidase gene. Proc Natl Acad Sci USA 99: 9043–9048
  31. ↑ Jung KH, Seo YS, Walia H, Cao P, Fukao T, Canlas PE, Amonpant F, Bailey-Serres J, Ronald PC (2010) The submergence tolerance regulator Sub1A mediates stress-responsive expression of AP2/ERF transcription factors. Plant Physiol 152: 1674–1692
  32. ↑ Cite error: Invalid <ref> tag; no text was provided for refs named ref276

Cite error: <ref> tag with name "ref27" defined in <references> is not used in prior text.

Structured Information

Gene Name

Os09g0457900

Description

Similar to AP2 domain containing protein RAP2.6 (Fragment)

Version

NM_001069906.1 GI:115479554 GeneID:4347266

Length

1100 bp

Definition

Oryza sativa Japonica Group Os09g0457900, 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 9

Location

Chromosome 9:17959811..17960910

Sequence Coding Region

17959874..17960698

Expression

GEO Profiles:Os09g0457900

Genome Context

<gbrowseImage1> name=NC_008402:17959811..17960910 source=RiceChromosome09 preset=GeneLocation </gbrowseImage1>

Gene Structure

<gbrowseImage2> name=NC_008402:17959811..17960910 source=RiceChromosome09 preset=GeneLocation </gbrowseImage2>

Coding Sequence

<cdnaseq>atgaccaagaaggtgataccggccatggcggcggcgaggcaggattcttgcaagaccaagcttgatgagcgtgggggtagtcatcaggctccgagctccgcgcggtggatctcgtccgagcaggagcacagcatcatcgtcgcggctctgcggtacgtggtgtccgggtgcaccacgccgccgccggagatcgtcacggtggcgtgcggggaggcgtgtgctctgtgcggcatcgacggctgtctcgggtgcgacttctttggggccgaggcggcggggaacgaggaggcggtaatggcgacggattatgctgctgctgctgctgcggccgcggtggcaggaggatcaggcgggaagagggttaggcggaggaggaagaagaacgtgtaccgcggcgtgcggcatcggccgtgggggaagtgggcagcggagatacgcgacccgcgccgcgcggtgcgcaagtggctcgggacgttcgacaccgccgaggaggccgccagggcgtacgaccgcgccgccctcgagttccgcggcgcgcgcgcgaagctcaacttcccgtgctccgagcctttgcccatgcccagccaaagaaacggcaatggcggcgatgctgtcacggcggcgacgacaacggcagagcagatgactccgactctgtcgccgtgcagcgcggatgccgaggagacgacgacgccggtggattggcagatgggcgcggacgaagccggcagcaaccagctctgggatggcttgcaggacctgatgaagctggatgaagcggacacctggttcccgccattttccggtgcagcgtctagtttttga</cdnaseq>

Protein Sequence

<aaseq>MTKKVIPAMAAARQDSCKTKLDERGGSHQAPSSARWISSEQEHS IIVAALRYVVSGCTTPPPEIVTVACGEACALCGIDGCLGCDFFGAEAAGNEEAVMATD YAAAAAAAAVAGGSGGKRVRRRRKKNVYRGVRHRPWGKWAAEIRDPRRAVRKWLGTFD TAEEAARAYDRAALEFRGARAKLNFPCSEPLPMPSQRNGNGGDAVTAATTTAEQMTPT LSPCSADAEETTTPVDWQMGADEAGSNQLWDGLQDLMKLDEADTWFPPFSGAASSF</aaseq>

Gene Sequence

<dnaseqindica>64..888#aacgacctcaagcacactactagctccggctcacttagctccctactgatcactggaacactcatgaccaagaaggtgataccggccatggcggcggcgaggcaggattcttgcaagaccaagcttgatgagcgtgggggtagtcatcaggctccgagctccgcgcggtggatctcgtccgagcaggagcacagcatcatcgtcgcggctctgcggtacgtggtgtccgggtgcaccacgccgccgccggagatcgtcacggtggcgtgcggggaggcgtgtgctctgtgcggcatcgacggctgtctcgggtgcgacttctttggggccgaggcggcggggaacgaggaggcggtaatggcgacggattatgctgctgctgctgctgcggccgcggtggcaggaggatcaggcgggaagagggttaggcggaggaggaagaagaacgtgtaccgcggcgtgcggcatcggccgtgggggaagtgggcagcggagatacgcgacccgcgccgcgcggtgcgcaagtggctcgggacgttcgacaccgccgaggaggccgccagggcgtacgaccgcgccgccctcgagttccgcggcgcgcgcgcgaagctcaacttcccgtgctccgagcctttgcccatgcccagccaaagaaacggcaatggcggcgatgctgtcacggcggcgacgacaacggcagagcagatgactccgactctgtcgccgtgcagcgcggatgccgaggagacgacgacgccggtggattggcagatgggcgcggacgaagccggcagcaaccagctctgggatggcttgcaggacctgatgaagctggatgaagcggacacctggttcccgccattttccggtgcagcgtctagtttttgagctagtgttattagatctcaaccgttggattagattactggaaggccattcatttattcgtttcattgtatagctaattagctatacttcattgtttgtgcagagcacgttttttaggagtaccgtcttatacatatttttttgtacagagtagaaagtaacaagaattttgtatattgatggaatagattatttccaattaatatttgcgc</dnaseqindica>

External Link(s)

NCBI Gene:Os09g0457900, RefSeq:Os09g0457900

Retrieved from "https://ngdc.cncb.ac.cn/ricewiki/index.php?title=Os09g0457900&oldid=178875"