Difference between revisions of "Os09g0104200"

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===Function===
 
===Function===
 
[[File: Shijc-Os09g0104200-Fig1.png|right|thumb|200px|'' Phenotypic characterization of vegetative and reproductive organs of osrad51d mutant and Ubi:RNAi-OsRAD51D transgenic rice plants. (from reference <ref name="ref1" />).'']]
 
[[File: Shijc-Os09g0104200-Fig1.png|right|thumb|200px|'' Phenotypic characterization of vegetative and reproductive organs of osrad51d mutant and Ubi:RNAi-OsRAD51D transgenic rice plants. (from reference <ref name="ref1" />).'']]
−
Osrad51d knock-out and RNAi-mediated knock-down transgenic (Ubi:RNAi-OsRAD51D) plants (Figure 2) revealed that suppression of OsRAD51D exerted negligible effects on the vegetative growth of rice plants (Figure 3a). In contrast, loss of OsRAD51D caused defects in reproductive growth. Homozygous mutant flowers displayed impaired development of pollen (Figure 3b–d) and Table 1), lemma and palea (Figure 4a,b), stamen, and carpel (Figure 4c and Table 2), which resulted in sterile flowers (Figure 4d). The abnormalities of heterozygous mutant and Ubi:RNAi-OsRAD51D plants were intermediate between wild type and homozygous mutant plants.
+
Osrad51d knock-out and RNAi-mediated knock-down transgenic (Ubi:RNAi-OsRAD51D) plants revealed that suppression of OsRAD51D exerted negligible effects on the vegetative growth of rice plants. In contrast, loss of OsRAD51D caused defects in reproductive growth. Homozygous mutant flowers displayed impaired development of pollen, lemma and palea, stamen, and carpel, which resulted in sterile flowers. The abnormalities of heterozygous mutant and Ubi:RNAi-OsRAD51D plants were intermediate between wild type and homozygous mutant plants.
 
[[File: Shijc-Os09g0104200-Fig2.png|right|thumb|200px|'' Cytogenetic analysis of root tips, PMCs, and pollen cells from wild type (WT) andosrad51d mutant rice plants. (from reference <ref name="ref1" />).'']]
 
[[File: Shijc-Os09g0104200-Fig2.png|right|thumb|200px|'' Cytogenetic analysis of root tips, PMCs, and pollen cells from wild type (WT) andosrad51d mutant rice plants. (from reference <ref name="ref1" />).'']]
−
Furthermore, previous studies in Arabidopsis indicate that AtRAD51B, AtRAD51D, and AtXRCC2 are not essential for meiosis <ref name="ref2" /><ref name="ref3" /><ref name="ref4" />(Bleuyard et al., 2005; Durrant et al., 2007; Da Ines et al., 2013). Triple, as well as single/double, mutations of AtRAD51B, AtRAD51D, and AtXRCC2 result in normal growth and fertility <ref name="ref5" />(Wang et al., 2013). These results are in sharp contrast with our results in that loss of OsRAD51D function caused severe errors in floral development and infertility (Figures 3 and 4). Cytogenetic analysis showed that osrad51d PMCs were unable to form normal homologous chromosome pairings and most mutant chromosomes were fragmented and scattered throughout the cells during meiosis I (Figure 5b). Due to these meiotic anomalies, homozygous osrad51d pollen cells contained numerous micro-nuclei and formed atypical tetrads, resulting in malfunctioning pollen (Figure 5c). Taken together, our results are consistent with the view that, unlike Arabidopsis AtRAD51D, OsRAD51D is critically involved in meiosis and is an essential factor for reproductive growth in rice plants. Nevertheless, it is still possible that absence of OsRAD51D could be affecting cellular stability of other OsRAD51 paralogs and thus producing an indirect effect on meiosis and floral development. A previous study of RAD51 in moss Physcomitrella patens suggested that there are fundamental differences in the use of the HR pathway in different plant species, and that an essential role of RAD51 in viability does not correlate with organism taxonomic complexity<ref name="ref6" />(Markmann-Mulisch et al., 2007).  
+
Furthermore, previous studies in Arabidopsis indicate that AtRAD51B, AtRAD51D, and AtXRCC2 are not essential for meiosis. Triple, as well as single/double, mutations of AtRAD51B, AtRAD51D, and AtXRCC2 result in normal growth and fertility <ref name="ref5" />. These results are in sharp contrast with our results in that loss of OsRAD51D function caused severe errors in floral development and infertility. Cytogenetic analysis showed that osrad51d PMCs were unable to form normal homologous chromosome pairings and most mutant chromosomes were fragmented and scattered throughout the cells during meiosis I. Due to these meiotic anomalies, homozygous osrad51d pollen cells contained numerous micro-nuclei and formed atypical tetrads, resulting in malfunctioning pollen. Taken together, our results are consistent with the view that, unlike Arabidopsis AtRAD51D, OsRAD51D is critically involved in meiosis and is an essential factor for reproductive growth in rice plants. Nevertheless, it is still possible that absence of OsRAD51D could be affecting cellular stability of other OsRAD51 paralogs and thus producing an indirect effect on meiosis and floral development. A previous study of RAD51 in moss Physcomitrella patens suggested that there are fundamental differences in the use of the HR pathway in different plant species, and that an essential role of RAD51 in viability does not correlate with organism taxonomic complexity<ref name="ref6" />.  
  
 
===Mutation===
 
===Mutation===
 
[[File: Shijc-Os09g0104200-Fig3.png|right|thumb|200px|'' Molecular characterization of T-DNA osrad51d knock-out mutant and Ubi:RNAi-OsRAD51D knock-down transgenic rice plants. (from reference <ref name="ref1" />).'']]
 
[[File: Shijc-Os09g0104200-Fig3.png|right|thumb|200px|'' Molecular characterization of T-DNA osrad51d knock-out mutant and Ubi:RNAi-OsRAD51D knock-down transgenic rice plants. (from reference <ref name="ref1" />).'']]
−
A single loss-of-function rice mutant line that contained a T-DNA insertion in the OsRAD51D gene was obtained (Yi and An, 2013; http://signal.salk.edu/cgi-bin/RiceGE). This mutant was named osrad51d. The T-DNA insertion was mapped to the third exon in OsRAD51D located on chromosome 9 (line PFG_4A-00300.R; Figure 2a). A homozygous line for the T-DNA insertion was identified by genomic PCR with FW2/RV3 and RB/RV3 primer sets (Figure 2b). Disruption of OsRAD51D by T-DNA insertion was further verified by RT-PCR. The results showed that the homozygous osrad51d mutant leaves contained undetectable amounts of all three OsRAD51D isoform transcripts under the experimental conditions, whereas the heterozygous osrad51d line possessed reduced levels (approximately 50%) of the transcripts (Figure 2c). Genomic Southern blot analysis indicated that the osrad51d mutant progeny contained a single T-DNA integration into the OsRAD51D gene (Figure 2d). Because only one allele of the osrad51d loss-of-function mutant plant was obtained, RNAi-mediated knock-down transgenic rice lines (Ubi:RNAi-OsRAD51D) were generated. RT-PCR revealed that the mRNA levels of OsRAD51D.1/OsRAD51D.3 and OsRAD51D.2 significantly decreased in the independent T2 Ubi:RNAi-OsRAD51D lines #1 and #2 (Figure 2c). Genomic Southern blotting demonstrated that these two RNAi-transgenic plants were independent lines (Figure 2d).
+
A single loss-of-function rice mutant line that contained a T-DNA insertion in the OsRAD51D gene was obtained (Yi and An, 2013; http://signal.salk.edu/cgi-bin/RiceGE). This mutant was named osrad51d. The T-DNA insertion was mapped to the third exon in OsRAD51D located on chromosome 9 (line PFG_4A-00300.R). A homozygous line for the T-DNA insertion was identified by genomic PCR with FW2/RV3 and RB/RV3 primer sets. Disruption of OsRAD51D by T-DNA insertion was further verified by RT-PCR. The results showed that the homozygous osrad51d mutant leaves contained undetectable amounts of all three OsRAD51D isoform transcripts under the experimental conditions, whereas the heterozygous osrad51d line possessed reduced levels (approximately 50%) of the transcripts. Genomic Southern blot analysis indicated that the osrad51d mutant progeny contained a single T-DNA integration into the OsRAD51D gene. Because only one allele of the osrad51d loss-of-function mutant plant was obtained, RNAi-mediated knock-down transgenic rice lines (Ubi:RNAi-OsRAD51D) were generated. RT-PCR revealed that the mRNA levels of OsRAD51D.1/OsRAD51D.3 and OsRAD51D.2 significantly decreased in the independent T2 Ubi:RNAi-OsRAD51D lines #1 and #2. Genomic Southern blotting demonstrated that these two RNAi-transgenic plants were independent lines .
  
 
===Expression===
 
===Expression===
 
[[File: Shijc-Os09g0104200-Fig4.png|right|thumb|200px|'' Identification and expression of OsRAD51D in rice. (from reference <ref name="ref1" />).'']]
 
[[File: Shijc-Os09g0104200-Fig4.png|right|thumb|200px|'' Identification and expression of OsRAD51D in rice. (from reference <ref name="ref1" />).'']]
−
OsRAD51D.1/OsRAD51D.3 and OsRAD51D.2 isoforms were expressed in all tissues examined in rice plants (Figure 1d). A FW1/RV1 primer set detected both OsRAD51D.1 and OsRAD51D.3 mRNAs due to their similar structure. Yeast two-hybrid assay showed that human RAD51B and RAD51D interacted in the presence of RAD51C (Schild et al., 2000). In vivo immuno-precipitation results indicated that human AD51 paralogs form the RAD51B-RAD51C-RAD51D-XRCC2 complex (Masson et al., 2001; Liu et al., 2002). Arabidopsis AtRAD51B and AtRAD51C also interact in yeast cells <ref name="ref7" />(Osakabe et al., 2005). Yeast two-hybrid studies showed that rice OsRAD51D.1 interacted with OsRAD51B and OsRAD51C, whereas OsRAD51D.2 and OsRAD51D.3 were unable to associate with other OsRAD51 paralogs (Figure 1e). In vitro pull-down assays also indicated that bacterially expressed OsRAD51D.1, but not OsRAD51D.2 and OsRAD51D.3, was associated with OsRAD51B and OsRAD51C (Figures 1f and S3). These results raise the possibility that the OsRAD51D.1 isoform is the major OsRAD51D protein in rice plants.
+
OsRAD51D.1/OsRAD51D.3 and OsRAD51D.2 isoforms were expressed in all tissues examined in rice plants. A FW1/RV1 primer set detected both OsRAD51D.1 and OsRAD51D.3 mRNAs due to their similar structure. Yeast two-hybrid assay showed that human RAD51B and RAD51D interacted in the presence of RAD51C. In vivo immuno-precipitation results indicated that human AD51 paralogs form the RAD51B-RAD51C-RAD51D-XRCC2 complex. Arabidopsis AtRAD51B and AtRAD51C also interact in yeast cells <ref name="ref7" />. Yeast two-hybrid studies showed that rice OsRAD51D.1 interacted with OsRAD51B and OsRAD51C, whereas OsRAD51D.2 and OsRAD51D.3 were unable to associate with other OsRAD51 paralogs. In vitro pull-down assays also indicated that bacterially expressed OsRAD51D.1, but not OsRAD51D.2 and OsRAD51D.3, was associated with OsRAD51B and OsRAD51C. These results raise the possibility that the OsRAD51D.1 isoform is the major OsRAD51D protein in rice plants.
  
 
===Splicing Variants===
 
===Splicing Variants===
−
There are five RAD51 paralogs, OsRAD51B, OsRAD51C, OsRAD51D, OsXRCC2, and OsXRCC3, in rice (Figure S1). The rice OsRAD51D gene is 5804 bp in length and composed of nine exons and eight introns (Figure 1a). Three splicing variants of OsRAD51D were predicted and termed OsRAD51D.1 (GenBank accession no. KJ472480), OsRAD51D.2 (GenBank accession no. KJ472481), and OsRAD51D.3 (GenBank accession no. KJ472482) (Figure 1b). The coding regions of OsRAD51D.1, OsRAD51D.2, and OsRAD51D.3 are 849 bp encoding 283 amino acids (30.3 kDa), 552 bp encoding 184 amino acids (19.8 kDa), and 768 bp encoding 256 amino acids (27.4 kDa), respectively (Figure 1c). All three deduced OsRAD51D proteins contained Walker A and Walker B motifs and a RecA domain in their central regions (Figure 1c). Walker A and Walker B motifs function as nucleotide binding domains <ref name="ref8" />(Hanson and Whiteheart, 2005), whereas the RecA motif is an ATP hydrolysis domain <ref name="ref9" />(Cox, 2007). Both Walker A and Walker B motifs are highly conserved in plant (rice, wheat, maize, and Arabidopsis) and human RAD51D homologs (Figure S2a,b). The RecA domain in OsRAD51D is 45–76% and 29% identical to those in plant and human RAD51D proteins, respectively.
+
There are five RAD51 paralogs, OsRAD51B, OsRAD51C, OsRAD51D, OsXRCC2, and OsXRCC3, in rice. The rice OsRAD51D gene is 5804 bp in length and composed of nine exons and eight introns. Three splicing variants of OsRAD51D were predicted and termed OsRAD51D.1 (GenBank accession no. KJ472480), OsRAD51D.2 (GenBank accession no. KJ472481), and OsRAD51D.3 (GenBank accession no. KJ472482). The coding regions of OsRAD51D.1, OsRAD51D.2, and OsRAD51D.3 are 849 bp encoding 283 amino acids (30.3 kDa), 552 bp encoding 184 amino acids (19.8 kDa), and 768 bp encoding 256 amino acids (27.4 kDa), respectively All three deduced OsRAD51D proteins contained Walker A and Walker B motifs and a RecA domain in their central regions. Walker A and Walker B motifs function as nucleotide binding domains <ref name="ref8" />, whereas the RecA motif is an ATP hydrolysis domain <ref name="ref9" />. Both Walker A and Walker B motifs are highly conserved in plant (rice, wheat, maize, and Arabidopsis) and human RAD51D homologs. The RecA domain in OsRAD51D is 45–76% and 29% identical to those in plant and human RAD51D proteins, respectively.
  
 
==Labs working on this gene==
 
==Labs working on this gene==

Revision as of 14:01, 27 December 2014

OsRAD51D plays a critical role in reproductive growth in rice. [1]

Annotated Information

Function

Phenotypic characterization of vegetative and reproductive organs of osrad51d mutant and Ubi:RNAi-OsRAD51D transgenic rice plants. (from reference [1]).

Osrad51d knock-out and RNAi-mediated knock-down transgenic (Ubi:RNAi-OsRAD51D) plants revealed that suppression of OsRAD51D exerted negligible effects on the vegetative growth of rice plants. In contrast, loss of OsRAD51D caused defects in reproductive growth. Homozygous mutant flowers displayed impaired development of pollen, lemma and palea, stamen, and carpel, which resulted in sterile flowers. The abnormalities of heterozygous mutant and Ubi:RNAi-OsRAD51D plants were intermediate between wild type and homozygous mutant plants.

Cytogenetic analysis of root tips, PMCs, and pollen cells from wild type (WT) andosrad51d mutant rice plants. (from reference [1]).

Furthermore, previous studies in Arabidopsis indicate that AtRAD51B, AtRAD51D, and AtXRCC2 are not essential for meiosis. Triple, as well as single/double, mutations of AtRAD51B, AtRAD51D, and AtXRCC2 result in normal growth and fertility [2]. These results are in sharp contrast with our results in that loss of OsRAD51D function caused severe errors in floral development and infertility. Cytogenetic analysis showed that osrad51d PMCs were unable to form normal homologous chromosome pairings and most mutant chromosomes were fragmented and scattered throughout the cells during meiosis I. Due to these meiotic anomalies, homozygous osrad51d pollen cells contained numerous micro-nuclei and formed atypical tetrads, resulting in malfunctioning pollen. Taken together, our results are consistent with the view that, unlike Arabidopsis AtRAD51D, OsRAD51D is critically involved in meiosis and is an essential factor for reproductive growth in rice plants. Nevertheless, it is still possible that absence of OsRAD51D could be affecting cellular stability of other OsRAD51 paralogs and thus producing an indirect effect on meiosis and floral development. A previous study of RAD51 in moss Physcomitrella patens suggested that there are fundamental differences in the use of the HR pathway in different plant species, and that an essential role of RAD51 in viability does not correlate with organism taxonomic complexity[3].

Mutation

Molecular characterization of T-DNA osrad51d knock-out mutant and Ubi:RNAi-OsRAD51D knock-down transgenic rice plants. (from reference [1]).

A single loss-of-function rice mutant line that contained a T-DNA insertion in the OsRAD51D gene was obtained (Yi and An, 2013; http://signal.salk.edu/cgi-bin/RiceGE). This mutant was named osrad51d. The T-DNA insertion was mapped to the third exon in OsRAD51D located on chromosome 9 (line PFG_4A-00300.R). A homozygous line for the T-DNA insertion was identified by genomic PCR with FW2/RV3 and RB/RV3 primer sets. Disruption of OsRAD51D by T-DNA insertion was further verified by RT-PCR. The results showed that the homozygous osrad51d mutant leaves contained undetectable amounts of all three OsRAD51D isoform transcripts under the experimental conditions, whereas the heterozygous osrad51d line possessed reduced levels (approximately 50%) of the transcripts. Genomic Southern blot analysis indicated that the osrad51d mutant progeny contained a single T-DNA integration into the OsRAD51D gene. Because only one allele of the osrad51d loss-of-function mutant plant was obtained, RNAi-mediated knock-down transgenic rice lines (Ubi:RNAi-OsRAD51D) were generated. RT-PCR revealed that the mRNA levels of OsRAD51D.1/OsRAD51D.3 and OsRAD51D.2 significantly decreased in the independent T2 Ubi:RNAi-OsRAD51D lines #1 and #2. Genomic Southern blotting demonstrated that these two RNAi-transgenic plants were independent lines .

Expression

Identification and expression of OsRAD51D in rice. (from reference [1]).

OsRAD51D.1/OsRAD51D.3 and OsRAD51D.2 isoforms were expressed in all tissues examined in rice plants. A FW1/RV1 primer set detected both OsRAD51D.1 and OsRAD51D.3 mRNAs due to their similar structure. Yeast two-hybrid assay showed that human RAD51B and RAD51D interacted in the presence of RAD51C. In vivo immuno-precipitation results indicated that human AD51 paralogs form the RAD51B-RAD51C-RAD51D-XRCC2 complex. Arabidopsis AtRAD51B and AtRAD51C also interact in yeast cells [4]. Yeast two-hybrid studies showed that rice OsRAD51D.1 interacted with OsRAD51B and OsRAD51C, whereas OsRAD51D.2 and OsRAD51D.3 were unable to associate with other OsRAD51 paralogs. In vitro pull-down assays also indicated that bacterially expressed OsRAD51D.1, but not OsRAD51D.2 and OsRAD51D.3, was associated with OsRAD51B and OsRAD51C. These results raise the possibility that the OsRAD51D.1 isoform is the major OsRAD51D protein in rice plants.

Splicing Variants

There are five RAD51 paralogs, OsRAD51B, OsRAD51C, OsRAD51D, OsXRCC2, and OsXRCC3, in rice. The rice OsRAD51D gene is 5804 bp in length and composed of nine exons and eight introns. Three splicing variants of OsRAD51D were predicted and termed OsRAD51D.1 (GenBank accession no. KJ472480), OsRAD51D.2 (GenBank accession no. KJ472481), and OsRAD51D.3 (GenBank accession no. KJ472482). The coding regions of OsRAD51D.1, OsRAD51D.2, and OsRAD51D.3 are 849 bp encoding 283 amino acids (30.3 kDa), 552 bp encoding 184 amino acids (19.8 kDa), and 768 bp encoding 256 amino acids (27.4 kDa), respectively All three deduced OsRAD51D proteins contained Walker A and Walker B motifs and a RecA domain in their central regions. Walker A and Walker B motifs function as nucleotide binding domains [5], whereas the RecA motif is an ATP hydrolysis domain [6]. Both Walker A and Walker B motifs are highly conserved in plant (rice, wheat, maize, and Arabidopsis) and human RAD51D homologs. The RecA domain in OsRAD51D is 45–76% and 29% identical to those in plant and human RAD51D proteins, respectively.

Labs working on this gene

1. Department of Systems Biology, College of Life Science and Biotechnology, Yonsei University, Seoul, Korea

References

  1. ↑ 1.0 1.1 1.2 1.3 1.4 Byun, M. Y. and Kim, W. T. (2014), Suppression of OsRAD51D results in defects in reproductive development in rice (Oryza sativa L.). The Plant Journal, 79: 256–269. doi: 10.1111/tpj.12558
  2. ↑ Wang, Y., Xiao, R., Wang, H., Cheng, Z., Li, W., Zhu, G., Wang, Y. and Ma, H. (2013) The Arabidopsis RAD51 paralogs RAD51B, RAD51D and XRCC2 play partially redundant roles in somatic DNA repair and gene regulation. New Phytol. 201, 292–304.
  3. ↑ Markmann-Mulisch, U., Wendeler, E., Zobell, O., Schween, G., Steinbiss, H.H. and Reissa, B. (2007) Differential requirements for RAD51 in Physcomitrella patens and Arabidopsis thaliana development and DNA damage repair. Plant Cell, 19, 3080–3089.
  4. ↑ Osakabe, K., Abe, K., Yamanouchi, H. et al. (2005) Arabidopsis Rad51B is important for double-strand DNA breaks repair in somatic cells. Plant Mol. Biol. 57, 819–833.
  5. ↑ Hanson, P.I. and Whiteheart, S.W. (2005) AAA+ proteins: have engine, will work. Nat. Rev. Mol. Cell Biol. 6, 519–529.
  6. ↑ Cox, M.M. (2007) Motoring along with the bacterial RecA protein. Nat. Rev. Mol. Cell Biol. 8, 127–138.

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Structured Information

Gene Name

Os09g0104200

Description

RecA bacterial DNA recombination family protein

Version

NM_001069094.2 GI:297609026 GeneID:4346376

Length

5790 bp

Definition

Oryza sativa Japonica Group Os09g0104200, 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 9:431724..437513

Sequence Coding Region

432061..432099,434262..434408,435614..435676

Expression

GEO Profiles:Os09g0104200

Genome Context

<gbrowseImage1> name=NC_008402:431724..437513 source=RiceChromosome09 preset=GeneLocation </gbrowseImage1>

Gene Structure

<gbrowseImage2> name=NC_008402:431724..437513 source=RiceChromosome09 preset=GeneLocation </gbrowseImage2>

Coding Sequence

<cdnaseq>atgatgatatcagtggctatgattctaaagaagctagcatatgagcataatctatctgttctggttactaaccatatggttgctgggaatggagctcccaagcctgctcttggtgagagctggaaaactgttccacatgtccgcttggtgatatctcgtgagcgtggtagcaaaatctgcgcagcaactgtgctgaagcacacactactggcttcgggccgtgttatgaaatttgccgtaccaagctga</cdnaseq>

Protein Sequence

<aaseq>MMISVAMILKKLAYEHNLSVLVTNHMVAGNGAPKPALGESWKTV PHVRLVISRERGSKICAATVLKHTLLASGRVMKFAVPS</aaseq>

Gene Sequence

<dnaseqindica>5415..5453#3106..3252#1838..1900#ttccccttctggctgtaatggcgatggcggcgacgtcggcgagcggcggcgccggctgcttccccaggcctcctccctccggcggcgaggttagtcctccgtcctcggctgcttgacggccatttcagttcatccccagcttttttattcctgcttcgaaatctcagcttaatcgggaacggaatgctgcacaaaaaccgtttgctcttgacgagataatgcgcatgcgattcacccgcagttggtttcacggcttcatactactgatcaatcgtattttctgtgtgtatgtggcatggatttgtgcttggggtacaggagaagaagcaagatgaggaggggcaatgctttctggatggaatggacttgctcaaggatgcgacggagaacaagcgcttcctccccacgggccttcaagggtactcctatcttcaatgtaccaatcatttcctttccctctctatctattgctattggccctcccccaacttgtttggatcataggattcccctaactgccttgctttagtttttttagtttttgggtaatttcactaccaaattgcagattttgacttcatcatgtcctcaattttccagcgtcgacgcgcttcttggaggcggcctgcgccaaggtcagctcactgaaataaccggccaatcgtcctccggtaaaacacaggtgaaaatgttaaccacttctatctttcttaacttattagacgcaatagatagatatacattttatatgcttactcctccggacaagattggcttgccaaaaacagtaggattttaaggtatacatcacaattcacaagcggcactgttgtttattctgtctatgtaactgcaggtctgtctttgttctgcttcacatgtggcagccaggcagttgggtgttgttatgtacttggatactagcaattccttttcccctagccgcattgctcgaatagttgatggattccccatctctttggtcagagaggtaggtaatcatacttgtacacatatttgaatactgctcctaccctgttacttggacttacctgaattgcttcctttgctgtggtgtcactaaatatagttgaagacatgtttctgctattagattactctaactagtaacacttgtttaacaacttgctatactacttgcttttttctaattaccacatttgacttgcagccaaagaatgtgcgactcgagagggtcatgagcagcatcatttgtaagtcagtctttgatatattcgatttgtttgaagtgctacatcaacttgaattgtcactgaagagcaaggtgagtgtgaagtgattgttcaatttgtgccttgaaaaactctgatgtgccacgtgctttttaggtaaacaatgggggtaacaagatatgcttgcttatcattgactcaatatcgtctatacttgctcccatcaacggtgggaagtatccacgaggtacttgcatagtctttgtgccctacttttattattcatagcataagttgcaccttgctattgaacaagtaaatcagtctcgataagcagggacgtcataagtggctgagatatactagatccgattagcaagttagaaatagcagaaaattagttcagtgagcacttaacctgcaattatacagcgagatattgatactttgatctggtcataaattcaatgtgcttctgttaattgttgaagttccaaaaatcatttattttctcacacaattacttcctgagagggtccttattaatttttaaagtttattatgaagtaaatttgttgtttccttttgtctctgcatgtcttattaacctaacagggcgatcgatgatgatatcagtggctatgattctaaagaagctagcatatgagcataatctatctgttctggtaattctactttagtctaatttgtgtgctaataatttttggcatttgcactgcacgctttgggttgagcatatctgcacaaggctttctaaaatattattctgactcatcaagctattgtatgcattatacttcttttaacacatcggttgtgcaataacacctttgacaatggtgtggctaagatcacattactacagcgttcacccctttcgattgaatttagattctcatcatcctttctcgtttaaccccactcaatccccatcctgtgaattacataactgtggcccccgcaccctcatggttaaggtcatgcaatgtcctgaactcctgattgtgcctactcctggtaggatgtatgctggccagagttgctttggtctccacaattttatattgcttgcacttccttacatacagagaggtgacccttaacctccaggttgcttcacatgaaataggcagataggaacaggtggggatacatttcaagatagaatctctagaggtcaggcgtttgagtactaaagaagaactgctgtggtcacgtgccctcactgacattttttacatgttggagctagttgtcagtccacgttatcacttaaggttatgatgggcaggttaaggatcttgtgtgaatcacttaatgcacattataccgttgtgcagattttcagtttggtcacctaaatatgccacttaaactaatgtagtaacccctagtgcaatttgctctaattaattgctgtacaaattcacttattcagcatgccatctttctcatgaacagattgggacatgtatctggaaatgcttcatttcctgctttacttctaagataagctattaaagtaaattgatcatgtactgttagttttagtgggtttgctcatgtccatctctgttattctgtggaaattttaacttctctatcttgagcacatattccctctactttctctagtttagttttcctggtttggaaaaaggcaccaaagcctctattacttggagcaacttttctttagttagttaccttgaagactgactaccaagtttcagaaggcttgctaaattagagaagtcaccataaggcttgctaacttgggcactatgcctgttgtgcttaaaatgtatgctgaaacaaaaatatggtcatcattttcatttatcggtttgcccccttgcattgactccaggttactaaccatatggttgctgggaatggagctcccaagcctgctcttggtgagagctggaaaactgttccacatgtccgcttggtgatatctcgtgagcgtggtagcaaaatctgcgcagcaactgtgctgaagcacacactactggtatttgcttaaacaattagattataagtttaaatgtcataatcatagcagcattgcttattgtctttctagaaattgcattaataatgtgttagctatcattttgttgaaaataagtggtgaagatattggcatgaggagaacaaaagtagtatactccctccatccataagtctaaggcatatttcatttctagtttttccaaataagatggcatatttgtagtcttcatgcatctaagacttaaatgaaggggttatcttttcgttttacggtgctaaattaaattgttgaccggaacccgagcaatcatcttaataatcattggttgcatgcatgcatatagttccttggtgtttgttatatgtgggatgagttaatttctccttggtcttgttgacaaaagaaatacccccttgttccacaaaatttacacctattacttttgtcaccaagaccaaggagaaattaactcatcttgcatgtaacaaagatcaagctgtgtacgcacacgtgcaaccaataagtattaagatgactcctcgggttccgatcaaacatttagtttacctctccatttaagtcttggatgcatagagactacaaatagaaacaacttatttggaaaaactcaaatatgaaataggtctaattcttgtggatggagggagtatgccttagacttttggatggagggagtaaacaattatatggtgccagtgcatacctctatatcctaatcctacatgttgcgtgttgttttcatgctatgctgcatttatatatttttgtatcatctttttttttctcactttttgtgcttttgaattgggctgtgaaaacaatagtggtaggtagatgccagatacatttttctcctatataaaaatttgagctggtagtgatggccatcagtcatcagtcatcccaagtcccctctaactcaagacaaactctgtaggatcctcatggaaaatttgcaaaagccacaaactcctttagactcttatttgtttgtaggaaatatgggtataccaccagtgtacctagccaatgaagctgcacatttaatgagtatttctatgttagatctgtcttatgatgaaatggaatggtctagtacctggcaatcctgcataaacgttctgagtactgcagagaggagactgagactaaagattatgttaattcattgcttaaagaaattacatggatcgattgcttctaatgtgggtggccctaaaaatctctaaaaataacttggtaactattaaaaaaagatacttgataactggaagataacttatctcttgactctcaagtctcaacatctgcatttgccgctcgctgtgtgtgtggccatcgcattcaaatcccatagcatctgaattgtgctggcgtctgagaatccatttgcctctggccacagtagacagttgtgacatgtaagaaaaaagtgatcatgtaaatgctatgccgctgagctcaaaattaatatgcaatatccatcataatttgttttagaaatacgcacttcaaaaataaattgttttcagttgacactttatatgagatggatgttgttctttcttagttttcttttcgttgtttttcttttatttattttgtaagcagacgcctgaggctcaaattccctttcttgggtgttcgaatttagggataaaggggtggaatatgggttagaatatcaggtggggagatgctaaccttccatctctttctaatgagttaggactgtctggccttattcatactccaatcctacttctttttgcacaaatacaaatctattggcatttctttgctctctaagaatcatgcaaggaaaagacggccattttgaatgatatgctgccctgccataaattaatctgcattatatcactgcaaaggaagttttttttaatctatttgcttcttttttgcagtaatgctcacaattgatgttttacactaaattacacccacttgacaagtttgtgcactaaattagacccactaaatgacacaagacgatgctaagtaccgtcatttcttccaatgtctggtagcaaaataaattgcatgattcttggtatcatctgtacatgcccaaaaacaaaaaagtgcgggcatgtcatctttgtttttcttcattgcaggcttcgggccgtgttatgaaatttgccgtaccaagctgaattcagcaacttgatgcttcatatataaccagaagggaactgatctttgtagttcaagctatagaaagaacttcacccgtaagggtaaccttttgatttaccttggctttatgttccaggccatttgtatattaatccaatggtttttgcatgttgcctgcgtccgtggtatcacgcttaactgatccatatatatagtttcttggtaaactgaatgtatcagaaacattcagttttcaagtttttgtttcttcaattattcactgtgaatactggaaaatggtagtttcccatcagtggataaaattgaatgaaattatatttgttccttgttttc</dnaseqindica>

External Link(s)

NCBI Gene:Os09g0104200, RefSeq:Os09g0104200

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