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| | ==Annotated Information== | | ==Annotated Information== |
| | ===Function=== | | ===Function=== |
| − | It reports here the molecular genetic characterization of dwarf27(d27), a classic rice mutant exhibiting increased tillers and reduced plant height.dwarf27 encodes a novel iron-containing protein that localizes in chloroplasts and is expressed mainly in vascular cells of shoots and roots. The phenotype of d27 is correlated with enhanced polar auxin transport. D27 is involved in the MAX/RMS/D pathway, in which D27 acts as a new member participating in the biosynthesis of strigolactones.Tillering in rice is one of the most important agronomic traits that determine grain yields and a model system for elucidating molecular mechanisms that regulate axillary buds (Wang and Li,2005). In this study, we characterize a rice dwarf 27 (d27) mutant that is defective in the outgrowth of axillary buds. Map-based cloning and in-depth analysis of D27 revealed that it encodes a novel chloroplast-located iron-containing protein. Our results demonstrate that D27 regulates tiller bud outgrowth through the MAX/RMS/D pathway and participates in the biosynthesis of strigolactones.
| + | Please input The Function of This Gene. |
| − | D27 Encodes a Novel Iron-Containing Protein Sequence analysis of 5 9 - and 3 9 -rapid amplification of cDNA ends (RACE) products indicated that the full length of D27 cDNA is 1254-bp long, with an ORF of 837 bp, a 217-bp 5 9 -untranslated region, and a 200-bp 3 9 -untranslated region (see Supplemental Figure 2 online). Sequence comparison between genomic DNA and cDNAs revealed that D27 is composed of seven exons that encodes a 278–amino acid polypeptide (Figure 2C; see Supple-
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| − | mental Figure 2 online). The 4-bp deletion in d27 results in a premature translational product (Figure 3A; see Supplemental Figure 2 online). The BLASTP (Altschul et al., 1997) analysis revealed that D27 shares no homology with any functionally identified protein and contains no conserved domain. However,analysis of multiple alignment against the National Center for Biotechnology Information database and The Institute for Genomic Research (TIGR) plant transcript assemblies showed thatD27 has homologies in many plant species, from lower plants to higher plants (see Supplemental Figure 3 online), suggesting that D27 may play a basic role in plants.Interestingly, when we tried to express and purify recombinant D27, we found that the bacterial cells expressing the maltose binding protein (MBP)-D27 fusion protein were strikingly brown in color, as was the purified MBP-D27 fusion protein (Figure 3B).This result suggested that D27 is very likely to have a cofactor. To explore this possibility, we analyzed the recombinant D27 with inductively coupled plasma mass spectrometry (ICP-MS) and found that the recombinant MBP-D27 protein contains ; 1.7mole of iron per mole of protein, in contrast with an extremely low level of iron bound to the C-terminal truncated polypeptide,
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| − | MBP-D27 1-187 (Figure 3C, Table 1), which is equivalent to the mutated form of D27. The binding of iron to D27 was further confirmed by characterizing the absorbtion spectrum of the MBP-D27 fusion protein, which showed a specific peak at 420nm, a characteristic for the presence of iron (Figure 3D). Fur-
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| − | thermore, when the recombinant MBP-D27 protein was treated with the reducing agent dithionite, the peak at 420 nm exhibited a dramatic decrease (Figure 3D), indicating that D27 is indeed an iron-containing protein. Moreover, the purified recombinant D27 protein contained no significant amount of other metals (Table 1), suggesting that the binding of iron to D27 is specific.Taken together, all these results indicate that D27 is an authentic iron-containing protein in plants.
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| − | Enhanced Polar Auxin Transport in d27
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| − | Shoot branching has been reported to be correlated to polar auxin transport (PAT) in Arabidopsis and pea (Morris, 1977;Beveridge et al., 2000; Bennett et al., 2006; Dai et al., 2006). We therefore investigated whether D27 is involved in PAT in rice. By comparing the basipetal and acropetal IAA transport in upper-
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| − | most internodes between the wild-type and d27 plants, we found that basipetal PAT in d27 was significantly elevated, whereas acropetal PAT of 3 H-IAA and basipetal transport of 3 H-IAA treated with the PAT inhibitor N-1-naphthylphtalamic acid (NPA) showed no significant difference between the wild-type and mutant plants (Figure 5A).To investigate whether the increased auxin transport is related to the d27 mutant phenotype, we further examined the effect of NPA on d27 seedlings in hydroponic culture. Two-week-old seedlings were treated with various concentrations of NPA. As shown in Figures 5B and 5C, the tiller number of d27 mutant plants was largely rescued when grown in the presence of 1.5 m M NPA for 5 weeks. Furthermore, when 2-week-old wild-type and d27 seedlings were treated with as low as 0.5 m M NPA, the outgrowth of tiller buds of the wild type was significantly pro-moted, which is consistent with the long-established concept that too little auxin transport also leads to an increased shoot branching (Chatfield et al., 2000). By contrast, the treatment of NPA showed a remarkable inhibition to the tiller outgrowth of the d27 seedlings, suggesting that the tillering phenotype of d27 may be correlated with an enhanced PAT.
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| − | D27 May Function through the MAX/RMS/D Pathway
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| − | In rice, one class of tillering dwarf mutants with an increased tiller number and reduced plant height has been previously reported (Kinoshita and Takahashi, 1991). Among them, d3, d17/htd1, and d10 are found to function as their orthologs of MAX2/RMS4,
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| − | MAX3/RMS5, and MAX4/RMS1/DAD1 in Arabidopsis, pea, and Petunia, respectively (Stirnberg et al., 2002; Sorefan et al., 2003;Booker et al., 2004; Foo et al., 2005; Ishikawa et al., 2005;Snowden et al., 2005; Johnson et al., 2006; Zou et al., 2006; Arite et al., 2007). The phenotype of d27 and the involvement of D27 in
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| − | PAT prompted us to test whether D27 is a new member of the MAX/RMS/D pathway in rice. We therefore generated a d27 d10 double mutant and compared the phenotypes of single and double mutants of d27, d10, and d27 d10. As shown in Figure 6,d10 exhibits similar phenotype to d27, but has more tillers and a more severe dwarf stature than d27 (Figures 6A and 6B). Phe-notypic analysis revealed that the d27 d10 double mutant showed similar tiller number and plant height to d10 (Figures
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| − | 6A to 6D). Further investigation on the responses of d27, d10,and d27 d10 to the treatment of different NPA concentrations indicated that the d27 d10 double mutant has a similar response to d10 (Figures 6E and 6F). These results strongly suggested that D27 participates in the MAX/RMS/D pathway.
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| − | D27 Participates in the Biosynthesis of Strigolactones
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| − | Recent studies have shown that the proposed novel hormonesthat inhibit plant branching and are derived from the MAX/RMS/Dpathway are strigolactones or their downstream metabolites. MAX1, MAX3/RMS5/D17, and MAX4/RMS1/D10 are involved in the biosynthesis of strigolactones, while MAX2/RMS4/D3 is involved in strigolactone signaling (Gomez-Roldan et al., 2008;Umehara et al., 2008). To further understand the role of D27 in the MAX/RMS/D pathway in rice, we investigated whether d27 is
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| − | deficient in strigolactone production or signaling. By applying 1.0m M GR24, a synthetic strigolactone analog that acts as native
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| − | strigolactones, to wild-type and d27 seedlings in a hydroponicculture, we found that the exogenous supplement of GR24 was able to fully inhibit tiller bud outgrowth of 3-week-old d27 seedlings (Figure 7A), and continuous treatment for 7 weeks fully restored the tillering dwarf phenotype of d27 (Figure 7B).We further analyzed and compared the strigolactones pro-duced in the root exudates of wild-type and d27 seedlings by liquid chromatography–quadruple/time-of-flight tandem mass spectrometry (LC/MS-MS). Our results clearly showed that2 9 -epi-5-deoxystrigol (epi-5DS), an identified strigolactone in the hydroponic culture media of rice seedlings, was produced in the wild-type cultivar Shiokari root exudates but was undetect-able in d27 (Figure 7C). Moreover, we performed a highly sensitive germination assay using Orobanche minor seeds to estimate the strigolactone production in d27 root exudates. In agreement with LC/MS-MS data, the germination-stimulating activity of d27 root exudates dramatically decreased in compar-ison with the wild type (Figure 7D; see Supplemental Figure 4 online). These results indicate that D27 is required for the production of strigolactones in rice.
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| − | D27 Is a Novel Iron binding Protein That Localizes in Chloroplasts
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| − | Although the bioinformatic analysis shows that D27 encodes a novel protein with no homology to any functionally known proteins, D27 homologs are found from algae (cyanophyta) to higher plants, but not in animals or fungi (see Supplemental Figure 3 online), suggesting that D27 may be a plant-specific protein. Analysis of the transient expression in rice protoplasts demonstrates that the D27 protein is localized in chloroplasts,similar to MAX3 and MAX4/D10 (Booker et al., 2004; Auldridge et al., 2006; Arite et al., 2007).Our data also showed that D27 is an iron-containing protein and that the truncated D27 loses the ability to bind iron (Figure 3C). Bacterial cells expressing truncated MBP-D27 1-187 arecolorless, and the purified protein does not have a 420-nm peak. By contrast, the full-length MBP-D27 fusion protein is brown and has the characteristic absorbance at 420 nm (Figures 3B and 3D), suggesting that the C terminus of D27 may contain an iron binding domain.
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| − | D27 Suppresses the Outgrowth of Rice Tiller Buds through
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| − | the MAX Pathway The development of shoot branching occurs in two steps, the initiation of the AM and the outgrowth of axillary buds. Unlike the elusive molecular mechanism that regulates AM initiation, the outgrowth of axillary buds is well understood due to the recent breakthrough in the MAX/RMS/D pathway. Studies on a number of mutants that display excess axillary branches, max in Arabi-dopsis (Stirnberg et al., 2002, 2007; Sorefan et al., 2003; Booker et al., 2004), rms in pea (Beveridge et al., 1994, 1996, 2000; Foo et al., 2001, 2005; Morris et al., 2001; Sorefan et al., 2003), and dad in petunia (Napoli, 1996; Snowden et al., 2005; Simons et al.,2007), have revealed the existence of a carotenoid-derived AM outgrowth regulating pathway. Although the outgrowth behav-iors between dicotyledonous and monocotyledonous axillary buds are different (for reviews, see McSteen and Leyser, 2005;Wang and Li, 2008), they appear to share a conserved branchingsignal pathway because orthologs of MAX2/RMS4, MAX3/RMS5, and MAX4/RMS1 have also been identified in rice; they are D3, HTD/D17, and D10, respectively (Ishikawa et al., 2005;Zou et al., 2006; Arite et al., 2007). Rice plants harboring individual loss-of-function mutations in these genes lead to more tillers and reduced plant height, a similar phenotype to those in Arabidopsis and pea, indicating their conserved func-tions in suppressing branch development in monocotyledonous plants.
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| − | The more tillers phenotype of d27 is ascribed to the extensive outgrowth of tiller buds, especially to the higher-order tiller buds,which are dormant in the wild-type plants (Figures 1A to 1F). Thecomparable morphology of d27 to that of the rice tillering dwarfmutant d3, htd1/d17, or d10 prompted us to test the hypothesis
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| − | that D27 is also involved in the MAX/RMS/D pathway. The analysis of the double mutant d27 d10 confirms the hypothesis.In the phenotypes tested, including tillering behavior, plant height, and response to NPA treatment, d27 d10 resembles d10 (Figure 6), suggesting that D27 may function the same as D10 in the MAX/RMS/D pathway. In agreement with this, D27 is expressed in roots and shoots, especially in the vasculature tissue of the plants (Figures 4B to 4J), an expression pattern similar to those of D10 and HTD1/D17. These results are con-sistent with a role in the biosynthesis of strigolactones. Further determination of strigolactone-related products in d27 and d10 will facilitate the understanding of the genetic relationship between D27 and D10.
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| − | D27 Is Required for the Biosynthesis of Strigolactones
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| − | The MAX/RMS/D pathway has been proven to interact with classic plant hormones auxin and cytokinin, but all the evidence obtained so far has demonstrated that the MAX/RMS/D-dependent branching signals are not attributed to any known hormones. Recent studies uncover the role of strigolactones or their metabolites acting as a new class of branching hor-mones, the signal derived from the MAX/RMS/D pathway (Gomez-Roldan et al., 2008; Umehara et al., 2008). Although previous studies have shown that strigolactones are derived from the carotenoid pathway and function as important signals
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| − | in establishing the interaction between plants and mycorrhizal fungi (Akiyama et al., 2005; Matusova et al., 2005), the biosyn- thetic and signaling pathways of strigolactones are poorly understood. MAX1, MAX3/RMS5/D17, and MAX4/RMS1/D10 are essential components for the biosynthesis of strigolac-tones, whereas MAX2/RMS4/D3 is involved in the perception of the signal. Our studies provide direct evidence that D27 is a new component of the MAX/RMS/D pathway and plays an essential role in biosynthesizing strigolactones. First, the d27 phenotype can be restored to the wild type upon supplemention with GR24 (Figures 7A and 7B). Second, the d27 root extract contains undetectable strigolactone, which is normally produced in the wild-type root extract (Figure 7C). Third, unlike the wild type, the d27 root exudates failed to stimulate the seed germination of O. minor (Figure 7D; see Supplemental Figure 4 online). Based on the findings that the D27 protein is localized in chloroplasts and contains iron and the fact that the complex structure of strigolactones should undergo a number of enzymatic reac- tions, including hydroxylation, epoxydation, oxidation, etc., to achieve its biosynthesis (Matusova et al., 2005), we hypothesize that D27 may participate in a redox reaction involved in the biosynthesis of strigolactones. Further biochemical experi-ments are required to confirm this possibility in the future.
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| | ===Expression=== | | ===Expression=== |
| − | In rice (Oryza sativa), the plastid-localized protein DWARF27 (OsD27) is necessary for SL biosynthesis, but the equivalent gene in Arabidopsis has not been identified.And DWARF27 (D27) have been identified with reduced strigolactone levels or strigolactone response.
| + | Please input The Function of This Gene. |
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| − | To isolate the D27 gene, we took a map-based cloning approach.D27 was primarily delimited in an interval of ; 3.0 centimorgans between the two molecular markers C189 and RM206 on the long arm of chromosome 11 (Figure 2A). To fine-map the D27 locus, we generated a large F2 mapping population derived from a cross between d27-ZF802 and its isogenetic lines ZF802. Of 21,000 F2 plants, 5200 mutant plants were used for fine-mapping, and the D27 locus was located between the two cleaved amplified polymorphic sequence (CAPS) markers P1 and P7 (Figure 2B). Screening with newly developed molecular markers P1 to P7 (see Supplemental Table 1 online), D27 was further placed within an 18-kb DNA region between the P3 and P6 markers and cosegregated with the P5 marker (Figure 2C). Within this region, there are two open reading frames (ORFs).Sequencing of these two ORFs of d27-ZF802 revealed a 4-bp deletion at the fourth exon of a putative gene, ORF LOC_Os11g37650, and this deletion results in a frame shift and generates a premature translation termination product (see Supplemental Figure 2 online).The identity of D27 was further confirmed by a genetic com-plementation test. The plasmid pD27C, containing a 9.25-kbgenomic DNA fragment consisting of a 2236-bp upstream se-quence, the entire D27 gene including seven exons and six introns, and a 2044-bp downstream region (Figure 2D), was introduced into a d27-Nipponbare mutant. All four transgenic lines of pD27C complement the d27 phenotype (Figure 2E).Therefore, ORF LOC_Os11g37650 is the rice D27 gene, and its 4-bp deletion is responsible for the altered phenotype of d27.
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| − | Expression Patterns of D27 and Subcellular Localization of the D27 Protein
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| − | Real-time PCR analysis revealed that the D27 expression level is high in axillary buds and young panicles, medium in shoot bases and culms, and low in roots, sheaths, and leaves (Figure 4A). The tissue-specific expression pattern of D27 was further examined using mRNA in situ hybridization. D27 was predominantly ex-
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| − | pressed in young leaves (Figure 4B), axillary buds (Figure 4C),inflorescence promodia (Figure 4D), lateral roots (Figure 4E), and crown roots (Figure 4H). Furthermore, D27 expression was detected in vascular cells at the shoot apex of the main stem and young leaves (Figures 4F and 4I), in the nodal vascular anastomosis (Figure 4G), and in large and small vascular bundles of the internodes (Figure 4J).To determine the subcellular localization of the D27 protein, we
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| − | performed a transient expression experiment of D27 in rice leaf protoplasts. The C terminus of D27 was fused with green fluorescent protein (GFP) under the control of cauliflower mosaic virus (CaMV) 35S promoter, and the construct was transferred into rice leaf protoplasts by the polyethylene glycol–mediated method. In contrast with the control, which was ubiquitous in protoplast cells, the D27-GFP fusion protein was predominantly localized in chloroplasts
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| | ===Evolution=== | | ===Evolution=== |
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| | ==Structured Information== | | ==Structured Information== |
| − | {{JaponicaGene|
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| − | GeneName = DWARF27|
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| − | Description = Putative uncharacterized protein|
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| − | Definition = Oryza sativa Japonica Group DWARF27, complete gene.|
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| − | tag =|
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| − | tid = DWARF27-TA|
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| − | pid = DWARF27-PA|
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| − | Version = FJ641055.1 GI:254946545 Gene ID: 3974662|
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| − | Length = 5390 bp|
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| − | Source = Oryza sativa Japonica Group (Japanese rice)
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| − | cultivar = Nipponbare
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| − | ORGANISM Oryza sativa Japonica Group
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| − | Eukaryota; Viridiplantae; Streptophyta; Embryophyta; Tracheophyta;
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| − | Spermatophyta; Magnoliophyta; Liliopsida; Poales; Poaceae; BEP
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| − | clade; Ehrhartoideae; Oryzeae; Oryza.
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| − |
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| − | Chromosome = [[:category:Indica Chromosome 1|Chromosome 1]]|
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| − | AP = Chromosome 11:222..5190|
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| − | CDS = 222..525,2082..2221,2438..2530,2891..2952,3514..3620,4977..5066,5150..5190|
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| − | GCID = <gbrowseImage1>
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| − | name=JaponicaChromosome11:222..5190
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| − | source=RiceJaponica11
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| − | preset=GeneLocation
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| − | </gbrowseImage1>|
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| − | GSID = <gbrowseImage2>
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| − | name=JaponicaChromosome11:222..5190
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| − | source=RiceJaponica11
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| − | preset=GeneLocation
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| − | </gbrowseImage2>|
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| − | CDNA = <cdnaseq>ATTCCCACCACAACCAAGATGCCCTCTCCATGCCATTTGGTCTCTTCTCTCTCCCTCCTTGCAAATTGCATGACCCTCTCTCTCTCTCTCCACTTCTCTCTATAAACCTTCCTCTCTCCCATAACTTCTTTCCATTTTCAACCTACAAATATACTAATCTCTCTCTAGCTAGTCTTCACCTACAAATCTCTCTCTCTCTCTCTCTCTCTCTCTCTCTCTTCATGGAGACCACCACGCTTGTGCTGCTTCTTCCTCATGGCGGCGCCGGCGGCGTACGGCCGGCGGCAGCGGCAACGGCGAAGCGAAGCTACGTGATGAGGAGGTGTTGCTCGACGGTGAGGGCGGTCATGGCGAGGCCGCAAGAGGCGCCGGCGTCGGCGCCGGCCAAGAAGACGGAGACGGCGGCGATGATGTCGACGGTGCAGACGGAGACGGCGGCGGCGCCGCCGGCGACGGTGTACCGGGACAGCTGGTTCGACAAGCTCGCCATTGGTTACCTGTCCAGGAACCTTCAAGAAGCTTCTGGTTCGTGCCTATAATCTACTGAATATATCTGATGTATTGTGTGCATGTACATGCAATCTTTCGGATGAATTAATGAATGATCAAATCAACCATGCATGCATATATGTGTGTATTTATGATGTTTTTGCCATGGCTGATTTGATTTTTTGAGTGGAAGAAATAACATATATATGGTGAATAAACCAATGTTCTTCAGTTATTTTTGTGACAATTAAATTATATGTGTATTCTACGATCATCCTTTTCTTCTTATCGTTTGGTTATTAATACCCTTCTCGTCCATGTGGAACATAAATTCTTTGATTTTTTTTATTACATGTTGCTTTGGCCAGCGTGGGTGTACTTGCAATTGCATGCTTTCTGTCATATTATAATTGAATTAGCTCTTGATTTTTACATGCAAAAGAATAATTTATTTATTGAGTTGCTGATGATCACGTAAGCCTATGCATGTGAGATGTGATGTGTTCTCCGGCCTAATTAAGTATCGACGTACTATATATATGCACGTACATCAAGTACGGCCGTATTTCATAATTTGATTATTGTTACTGTACCAGACAAACCTAGTGCTTAATTATTGACATATACCCAACATCGATTAAAGTCAGTGTGACACAATTGGCAGAAGAAGATAACAAAAATCATAGAAGACATATATACCACAACAGTAAAAATTACAGTGCAAAAAGGGATACCAGAAGCTTCTTAATTAGGTGAATGGTCACATTTAGTAGTAGTAATACACAGTAATAACACATTATTTGACGAATAAATTAAGCGGAAGTGCATGATCAGCAGCATATAAGCAAGCAACTACGAGAGGTCAACCGTTGATCCGTGAAAATGTGAGGTTCACATTCTAGGAATCTTTTAGTGCTTGAATCTAATCCTTTCGTTTGTTTTCCCGTGGACTTTTTACTTTGATCGAATAAACTTATCTGAATAATGCGAAGGTCAAACATCACATGTACGTGCCCGGTTCTTCTTCTAGAAATTCAGCCGCTTACTATAATCGATGCTGGTTTTTGAATTTTTTTAAAGACATTTATTTTGGAACAACTTAAAAAAAGTGCACAAACCAGGGTTTACCAAACCGTTTGAAGTGAGGTTACTGCCACTCCATGGTTTGGCAATTATCACGAGGTGCGTCGTGGTTCCAAAAACTAATAAGGTTATCGTATGTGATAACCGCGATAACTGACGGTTTTGTAAACCCTGCACAGACAACATAATTATATTTACAGGACTGTTTGGACTTCATTAACACTGTAGCATCCACTACATCCGTCACCTCCTTGGCTCTCTTACATGTTTAGCAAAGTCAATCATACAGACAAACCAGAAAAGTTCCCAACAGACACAGAATGCTACTTTTCTAGTATGGGTAATTAAGCATATAAGAGAAACTTGAAAGCATACAGGAAACCGATGTTCACTAGTCCTGAGTTTTTTTTTGGTGTTGTTCAGGTGCAAATTAACTGCAAAGGTTGATTCCCTGAAAGGAACTTGTAGCACTTGCTGACTGACACGAACATGCATGCAATGTAATGCAGGGCTAAAGAATGAAAAGGATGGCTACGAGAGCCTGATAGATGCCGCCCTAGCCATCTCAAGAATCTTCAGTCTGGATAAACAAAGCGAGATTGTGACCCAAGCTCTTGAAAGAGCACTTCCAAGCTACATCCTCACAATGGTAAGTACCATAATCCATGACAATTGGCAATCATGTATGAATTATTGAAATTTAAAAAACCTAAAACTTTTTTTTATTTGAACAGGAAGTAAAATTCAGCATTTATTCTCTCTTTTTTTTTTGAGGAAAATACAGAACTTCTGTAAAGGGAGGAAACTTTAAAAGTTCTCTTCTTTTATTTCTTACCTCAACTTTGATTGAGAATTCTGTCGGCAGATCAAGGTGATGATGCCACCTTCAAGATTTTCCAGGGAGTACTTTGCTGCATTCACCACGATATTTTTTCCTTGGTTGGTTGGGCCGTGTGAGGTATATATTACATACACAGTTCCTCCTTTTGTTACTTCAGTTCAGAAAGGAATAGCTGCCTGATACCTGATTAGTGCGTCTTCACAAGAATGAATTCATGATTGTGCCTCTGCTGAAGGTAGCCCTGCAGAATGAATTGATACGAAGCCACACGTTAATTTGAGAAATATTGCTACAGAAGATCTTAAAATGCTGTTGAATGTAGGTTGCGACATATAATATCTCTTTGTTTTCAGGACATTACATTGTTTAAAGTAGTTGTACGAAGCATTTGTGGTGCAAATATCATATAAAGTATGGAATAAATGCCCTTGTATGGATAACTTGTCTTCTGAGTGATCATTCTGTCATTGAACAAAGGTTATGGAATCTGAAGTTGAAGGAAGGAAAGAGAAAAACGTGGTATATATCCCCAAATGCAGGTAATTCAATCATCATCAAGAAATTGTTTCACAAGTTTGACTAGGAGGAACATTTGTAAAGAAAATTTCTTCAACTGCTGGGTATCACGTAACATATCGGTCAAACGGTTTCCTACTCCCAGCTCTTCATAGGCAGACTACAAAAATTTTCAGGTCTATAAGTAGTAACCTGATGATTTGCAATTAACATATGACTCTGCAGCCATCAATTAGTTCTGAATGAATTGTTGCAGTATGTCAACTCTTACTCAAGTTAATCCACTCATGAAAAACAAATTGAAGAACTATGGCTGGGACACAAGTTGTATTGTCAACGTCGTAATTGTGGCAACTGACAATCCTACTCACACTCCACAAGCTGCTTATTATCAATTCTACCCACGTTAGACAAACTACCATGATTAACACTGGGTTAAGAAAAAAATAGAATTTTAAGTGCAGATATATCAATAAATACTGAAGAACCACCTTAAAAATACATAAATACTGAAGAACAGGATTTCTACTGGTGTTGCTGTTTTTCTTTTCTTTTTTTTTAATCTCCAATCCTATTTGTGGCAGATTTCTGGAAAGTACAAATTGTGTTGGTATGTGCACAAACCTTTGCAAGATTCCATGCCAGAAGTTCATCCAAGATTCACTTGGCATGAAGGTCTACATGTCTCCCAGTAAGCTTCCTCTGCTCTGATTCATCAGGAAAGCCTGAATATCAGTGGGTAAACACAACAACTTAAATATAATTTGGTAGAATAAGGTAAAAATAATCAAACTCCATTCCTTTGCATAAAGTTGGCAAGAAACTAATAATGGATTGTCATATCAACATCATCTCAAAGTTGAAAATTTGGGTTGAAACCTGGTGTGCCTCTAACTTCTCAGGACATTGACCTAATTTTACTATAGCGGATCATTATACGGACTGCCGCTAAAACATAAGTGAACTCAAAATATCAACTTTCGACTTTAAGCAGAATTAGTGAAGTAAAGTTGAATCAATGTAATAGGTCCATAGACCACCAACGGTTTAGAGCTATGAGAAATACAGAACTTAATTTGACATATTAATTTCATGCCACCTCTACCTACCGACAATCTGAGGACTGTGCCCTTTTTGGGTATGGGAACCACTGGCCACCATTCAGGAAACAGAAAGGGCTGAAGTGGGGCCAACATGTTTGCAGGTTACTACCTCCGTTTTTTAATTTATAACGTCGATGACTTTTTAGATATATATGATCATTCGTCTTATTCAAAAAAAAATGCAATTATCATTTATTTTATTGTGACTTAATTTATCATCAAATGTTCTTTAAGCATGACTTAAATTTTTTTTATATTTGCACAATAATTTTGAATAAGATGAATGGTCAAATGTTTGTCAAAAAGTCAACAACGTCATACATTAAAAAACGGAGGGAGTACAAAGCAATCACAAGCAAGTTGTACAGAAGAAGAAAGTGCTAGCCCCTACTTGTCATTCTTAGTGGGAAACGGCAAATTGCATGGACAAGTTGTGGAACAGTAGTGTGAAATCCAGAAATCCACAATGTAGTGCTGTGAAAATACAGTACAAGCACCTTCACATGGCAGTGACGAGCCAGCATGTCTCGATAATCAGAGCAATATGGTTCAATTTCACTTATGAGAAGCACTAGTTTTGAATGTTAGTCTTCTATGGTGATACACTGAATATTTCAACATCAATCATTTTTTCTAGGTCAAAAGATACTGATATTTTATGACATCACCCCACAGAATCAGATATCAACCTCTGTTGGAATTTGCAAGGCTCTCTTTCCTTTTTATCCTGTCGAGTGGTTAGATCTAGGAGTAAGCAATGAATGTTATTTACTGCCAGTGCTACTAAATAGTCCTATACAGCTTATTTCCAATTAAGCAAGAAGGTTTTTTCCAATACTAGGTCAATAAAAACGAGAATAAAGCAGAGTCCCTAAAGATAATAGTGAATGTATCAAGAGAAATGTAACATAATTACTTACATCAACTATATATTTATTCTTCAGATTTTGAAGACATGAGCTGTGAGATGATATTTGGACAGCAACCTCCTGAAGATGACCCTGCATTGAAGCAGCCATGCTTCCGGACAAAATGTAAGGAATTCCGCACTGAGGCACCTTCATGGATTCAATCAACACTCCTCTATATTCATACTTTTTACTTCGTGTATATGCAGGCGTCGCAAAGCAGAATCATGGTGTGAATTGCTCCATCTGATCTGAAAGAAATTATCAATAGATAGATTTCAAATCAGTAAAATGCCTTAAGCTCCATTTCCTTTATTCCTTTGGAAAAAAAATTAGCACCATCATTGTTTTTGCCCACAACACCAGCATGTTTGGAACATACACTCTTCATTGTAATCCAAAAGTAATCTAAGAGGAAATGAAAGGCCCAACAGTAACCATTTTAAGGTA</cdnaseq>|
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| − | AA =<aaseq>METTTLVLLLPHGGAGGVRPAAAATAKRSYVMRRCCSTVRAVMARPQEAPASAPAKKTETAAMMSTVQTETAAAPPATVYRDSWFDKLAIGYLSRNLQEASGLKNEKDGYESLIDAALAISRIFSLDKQSEIVTQALERALPSYILTMIKVMMPPSRFSREYFAAFTTIFFPWLVGPCEVMESEVEGRKEKNVVYIPKCRFLESTNCVGMCTNLCKIPCQKFIQDSLGMKVYMSPNFEDMSCEMIFGQQPPEDDPALKQPCFRTKCVAKQNHGVNCSI</aaseq>|
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| − | DNA =
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| − | <dnaseqindica>1..304#1861..2000#2217..2309#2670..2731#3293..3399#4756..4845#4929..4969#ATTCCCACCACAACCAAGATGCCCTCTCCATGCCATTTGGTCTCTTCTCTCTCCCTCCTTGCAAATTGCATGACCCTCTCTCTCTCTCTCCACTTCTCTCTATAAACCTTCCTCTCTCCCATAACTTCTTTCCATTTTCAACCTACAAATATACTAATCTCTCTCTAGCTAGTCTTCACCTACAAATCTCTCTCTCTCTCTCTCTCTCTCTCTCTCTCTTCATGGAGACCACCACGCTTGTGCTGCTTCTTCCTCATGGCGGCGCCGGCGGCGTACGGCCGGCGGCAGCGGCAACGGCGAAGCGAAGCTACGTGATGAGGAGGTGTTGCTCGACGGTGAGGGCGGTCATGGCGAGGCCGCAAGAGGCGCCGGCGTCGGCGCCGGCCAAGAAGACGGAGACGGCGGCGATGATGTCGACGGTGCAGACGGAGACGGCGGCGGCGCCGCCGGCGACGGTGTACCGGGACAGCTGGTTCGACAAGCTCGCCATTGGTTACCTGTCCAGGAACCTTCAAGAAGCTTCTGGTTCGTGCCTATAATCTACTGAATATATCTGATGTATTGTGTGCATGTACATGCAATCTTTCGGATGAATTAATGAATGATCAAATCAACCATGCATGCATATATGTGTGTATTTATGATGTTTTTGCCATGGCTGATTTGATTTTTTGAGTGGAAGAAATAACATATATATGGTGAATAAACCAATGTTCTTCAGTTATTTTTGTGACAATTAAATTATATGTGTATTCTACGATCATCCTTTTCTTCTTATCGTTTGGTTATTAATACCCTTCTCGTCCATGTGGAACATAAATTCTTTGATTTTTTTTATTACATGTTGCTTTGGCCAGCGTGGGTGTACTTGCAATTGCATGCTTTCTGTCATATTATAATTGAATTAGCTCTTGATTTTTACATGCAAAAGAATAATTTATTTATTGAGTTGCTGATGATCACGTAAGCCTATGCATGTGAGATGTGATGTGTTCTCCGGCCTAATTAAGTATCGACGTACTATATATATGCACGTACATCAAGTACGGCCGTATTTCATAATTTGATTATTGTTACTGTACCAGACAAACCTAGTGCTTAATTATTGACATATACCCAACATCGATTAAAGTCAGTGTGACACAATTGGCAGAAGAAGATAACAAAAATCATAGAAGACATATATACCACAACAGTAAAAATTACAGTGCAAAAAGGGATACCAGAAGCTTCTTAATTAGGTGAATGGTCACATTTAGTAGTAGTAATACACAGTAATAACACATTATTTGACGAATAAATTAAGCGGAAGTGCATGATCAGCAGCATATAAGCAAGCAACTACGAGAGGTCAACCGTTGATCCGTGAAAATGTGAGGTTCACATTCTAGGAATCTTTTAGTGCTTGAATCTAATCCTTTCGTTTGTTTTCCCGTGGACTTTTTACTTTGATCGAATAAACTTATCTGAATAATGCGAAGGTCAAACATCACATGTACGTGCCCGGTTCTTCTTCTAGAAATTCAGCCGCTTACTATAATCGATGCTGGTTTTTGAATTTTTTTAAAGACATTTATTTTGGAACAACTTAAAAAAAGTGCACAAACCAGGGTTTACCAAACCGTTTGAAGTGAGGTTACTGCCACTCCATGGTTTGGCAATTATCACGAGGTGCGTCGTGGTTCCAAAAACTAATAAGGTTATCGTATGTGATAACCGCGATAACTGACGGTTTTGTAAACCCTGCACAGACAACATAATTATATTTACAGGACTGTTTGGACTTCATTAACACTGTAGCATCCACTACATCCGTCACCTCCTTGGCTCTCTTACATGTTTAGCAAAGTCAATCATACAGACAAACCAGAAAAGTTCCCAACAGACACAGAATGCTACTTTTCTAGTATGGGTAATTAAGCATATAAGAGAAACTTGAAAGCATACAGGAAACCGATGTTCACTAGTCCTGAGTTTTTTTTTGGTGTTGTTCAGGTGCAAATTAACTGCAAAGGTTGATTCCCTGAAAGGAACTTGTAGCACTTGCTGACTGACACGAACATGCATGCAATGTAATGCAGGGCTAAAGAATGAAAAGGATGGCTACGAGAGCCTGATAGATGCCGCCCTAGCCATCTCAAGAATCTTCAGTCTGGATAAACAAAGCGAGATTGTGACCCAAGCTCTTGAAAGAGCACTTCCAAGCTACATCCTCACAATGGTAAGTACCATAATCCATGACAATTGGCAATCATGTATGAATTATTGAAATTTAAAAAACCTAAAACTTTTTTTTATTTGAACAGGAAGTAAAATTCAGCATTTATTCTCTCTTTTTTTTTTGAGGAAAATACAGAACTTCTGTAAAGGGAGGAAACTTTAAAAGTTCTCTTCTTTTATTTCTTACCTCAACTTTGATTGAGAATTCTGTCGGCAGATCAAGGTGATGATGCCACCTTCAAGATTTTCCAGGGAGTACTTTGCTGCATTCACCACGATATTTTTTCCTTGGTTGGTTGGGCCGTGTGAGGTATATATTACATACACAGTTCCTCCTTTTGTTACTTCAGTTCAGAAAGGAATAGCTGCCTGATACCTGATTAGTGCGTCTTCACAAGAATGAATTCATGATTGTGCCTCTGCTGAAGGTAGCCCTGCAGAATGAATTGATACGAAGCCACACGTTAATTTGAGAAATATTGCTACAGAAGATCTTAAAATGCTGTTGAATGTAGGTTGCGACATATAATATCTCTTTGTTTTCAGGACATTACATTGTTTAAAGTAGTTGTACGAAGCATTTGTGGTGCAAATATCATATAAAGTATGGAATAAATGCCCTTGTATGGATAACTTGTCTTCTGAGTGATCATTCTGTCATTGAACAAAGGTTATGGAATCTGAAGTTGAAGGAAGGAAAGAGAAAAACGTGGTATATATCCCCAAATGCAGGTAATTCAATCATCATCAAGAAATTGTTTCACAAGTTTGACTAGGAGGAACATTTGTAAAGAAAATTTCTTCAACTGCTGGGTATCACGTAACATATCGGTCAAACGGTTTCCTACTCCCAGCTCTTCATAGGCAGACTACAAAAATTTTCAGGTCTATAAGTAGTAACCTGATGATTTGCAATTAACATATGACTCTGCAGCCATCAATTAGTTCTGAATGAATTGTTGCAGTATGTCAACTCTTACTCAAGTTAATCCACTCATGAAAAACAAATTGAAGAACTATGGCTGGGACACAAGTTGTATTGTCAACGTCGTAATTGTGGCAACTGACAATCCTACTCACACTCCACAAGCTGCTTATTATCAATTCTACCCACGTTAGACAAACTACCATGATTAACACTGGGTTAAGAAAAAAATAGAATTTTAAGTGCAGATATATCAATAAATACTGAAGAACCACCTTAAAAATACATAAATACTGAAGAACAGGATTTCTACTGGTGTTGCTGTTTTTCTTTTCTTTTTTTTTAATCTCCAATCCTATTTGTGGCAGATTTCTGGAAAGTACAAATTGTGTTGGTATGTGCACAAACCTTTGCAAGATTCCATGCCAGAAGTTCATCCAAGATTCACTTGGCATGAAGGTCTACATGTCTCCCAGTAAGCTTCCTCTGCTCTGATTCATCAGGAAAGCCTGAATATCAGTGGGTAAACACAACAACTTAAATATAATTTGGTAGAATAAGGTAAAAATAATCAAACTCCATTCCTTTGCATAAAGTTGGCAAGAAACTAATAATGGATTGTCATATCAACATCATCTCAAAGTTGAAAATTTGGGTTGAAACCTGGTGTGCCTCTAACTTCTCAGGACATTGACCTAATTTTACTATAGCGGATCATTATACGGACTGCCGCTAAAACATAAGTGAACTCAAAATATCAACTTTCGACTTTAAGCAGAATTAGTGAAGTAAAGTTGAATCAATGTAATAGGTCCATAGACCACCAACGGTTTAGAGCTATGAGAAATACAGAACTTAATTTGACATATTAATTTCATGCCACCTCTACCTACCGACAATCTGAGGACTGTGCCCTTTTTGGGTATGGGAACCACTGGCCACCATTCAGGAAACAGAAAGGGCTGAAGTGGGGCCAACATGTTTGCAGGTTACTACCTCCGTTTTTTAATTTATAACGTCGATGACTTTTTAGATATATATGATCATTCGTCTTATTCAAAAAAAAATGCAATTATCATTTATTTTATTGTGACTTAATTTATCATCAAATGTTCTTTAAGCATGACTTAAATTTTTTTTATATTTGCACAATAATTTTGAATAAGATGAATGGTCAAATGTTTGTCAAAAAGTCAACAACGTCATACATTAAAAAACGGAGGGAGTACAAAGCAATCACAAGCAAGTTGTACAGAAGAAGAAAGTGCTAGCCCCTACTTGTCATTCTTAGTGGGAAACGGCAAATTGCATGGACAAGTTGTGGAACAGTAGTGTGAAATCCAGAAATCCACAATGTAGTGCTGTGAAAATACAGTACAAGCACCTTCACATGGCAGTGACGAGCCAGCATGTCTCGATAATCAGAGCAATATGGTTCAATTTCACTTATGAGAAGCACTAGTTTTGAATGTTAGTCTTCTATGGTGATACACTGAATATTTCAACATCAATCATTTTTTCTAGGTCAAAAGATACTGATATTTTATGACATCACCCCACAGAATCAGATATCAACCTCTGTTGGAATTTGCAAGGCTCTCTTTCCTTTTTATCCTGTCGAGTGGTTAGATCTAGGAGTAAGCAATGAATGTTATTTACTGCCAGTGCTACTAAATAGTCCTATACAGCTTATTTCCAATTAAGCAAGAAGGTTTTTTCCAATACTAGGTCAATAAAAACGAGAATAAAGCAGAGTCCCTAAAGATAATAGTGAATGTATCAAGAGAAATGTAACATAATTACTTACATCAACTATATATTTATTCTTCAGATTTTGAAGACATGAGCTGTGAGATGATATTTGGACAGCAACCTCCTGAAGATGACCCTGCATTGAAGCAGCCATGCTTCCGGACAAAATGTAAGGAATTCCGCACTGAGGCACCTTCATGGATTCAATCAACACTCCTCTATATTCATACTTTTTACTTCGTGTATATGCAGGCGTCGCAAAGCAGAATCATGGTGTGAATTGCTCCATCTGATCTGAAAGAAATTATCAATAGATAGATTTCAAATCAGTAAAATGCCTTAAGCTCCATTTCCTTTATTCCTTTGGAAAAAAAATTAGCACCATCATTGTTTTTGCCCACAACACCAGCATGTTTGGAACATACACTCTTCATTGTAATCCAAAAGTAATCTAAGAGGAAATGAAAGGCCCAACAGTAACCATTTTAAGGTA</dnaseqindica>|
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| − | Link = [http://www.ncbi.nlm.nih.gov/nuccore/254946545?report=genbank]|
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| − | }}
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| | [[Category:Genes]] | | [[Category:Genes]] |
| | [[Category: Japonica DNA]] | | [[Category: Japonica DNA]] |
