Os04g0645100

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Annotated Information

Function

Rice (Oryza sativa) endosperm accumulates a massive amount of storage starch and storage proteins during seed development. However, little is known about the regulatory system involved in the production of storage substances. The rice flo2 mutation resulted in reduced grain size and starch quality. Map-based cloning identified FLOURY ENDOSPERM2 (FLO2), a member of a novel gene family conserved in plants, as the gene responsible for the rice flo2 mutation. FLO2 harbors a tetratricopeptide repeat motif, considered to mediate a protein-protein interactions. FLO2 was abundantly expressed in developing seeds coincident with production of storage starch and protein, as well as in leaves, while abundant expression of its homologs was observed only in leaves. The flo2 mutation decreased expression of genes involved in production of storage starch and storage proteins in the endosperm. Differences between cultivars in their responsiveness of FLO2 expression during high-temperature stress indicated that FLO2 may be involved in heat tolerance during seed development. Overexpression of FLO2 enlarged the size of grains significantly. These results suggest that FLO2 plays a pivotal regulatory role in rice grain size and starch quality by affecting storage substance accumulation in the endosperm.

Expression and Mutation

The transcript of FLO2 is abundant in immature seeds and mature leaves, but not abundant in panicles before heading, stems, and roots.

Expression of the genes involved in storage starch production was greatly reduced in the flo2 mutant (Figure 1). The molecular structure of the storage starch accumulated in flo2 endosperm was also quite different from that of the wild type. Phenotypes similar to those of the flo2 mutant are found in several mutants of the enzymes involved in starch biosynthesis, but many differences are evident in their features. For example, the amylopectin properties of the flo2 grain largely resemble those of the mutant lacking BEI, although the endosperm of this mutant seed exhibits a normal phenotype and contains the same amount of starch as the wild type. The flo2 mutant showed a decrease of BEI protein (Figure 2), suggesting a reduction of BEI activity. Therefore, it is reasonable to state that the reduced BEI activity in the flo2 mutant greatly contributed to the aberrant structure of storage starch. The mutant of BEIIb also produces a white-core endosperm, but its amylopectin structure differs greatly from that of the flo2 mutant where the long side chains contained in its amylopectin are highly enriched in the BEIIb mutant. Amylose is mainly synthesized by the activity of GBSS, suggesting that the decrease of amylose content could be attributed to the reduced expression of GBSSI in the flo2 mutant. It has been shown that AGPase activity strongly affects the yield of grain starch. ISA1 and PUL, both of which have activities as starch debranching enzymes, are essential for the formation of the highly organized clustered structure of amylopectin. Therefore, aberrant features of storage starch in the flo2 endosperm are ascribed to the combined effect of reduced expression of these genes.

Figure 1. Phylogenetic Tree of the Representative FLO2 Homologs.(from reference) .

Figure 2. Phylogenetic Tree of the Representative FLO2 Homologs.(from reference) .

The flo2 mutant also showed significantly decreased expression of storage protein genes. In addition, decreased expression was detected for several enzymes involved in metabolism (Figure 3). In particular, Susy is considered a key enzyme for the production of storage starch as well as cellulose synthesis. PPDK has been reported as the gene responsible for the rice flo4 mutant, which has a similar appearance to the flo2 mutant. PGI is important for both glycolysis and gluconeogenesis by catalyzing the reversible isomerization of glucose-6-phosphate to fructose-6-phosphate. Alanine aminotransferase is involved in the efficiency of nitrogen uptake and grain weight increase. These facts suggest that FLO2 widely affects the processes involved in storage substance accumulation in rice endosperm.

There are several reports on the regulatory factors involved in the production of storage substances in endosperm. It has been suggested that there are some regulatory factors that directly or indirectly control the genes involved in the individual enzymes or genes encoding the storage proteins. In the case of direct regulation of gene expression, specific regulatory factors, which interact with the specific nucleotide sequences in the promoter region of the target gene, are proposed.

FLO2 has a TPR motif composed of tandem repeats of 34 amino acid residues. The TPR motif adopts a helix-turn-helix structure and may mediate protein–protein interactions. We detected two candidate proteins, an LEA protein and a bHLH protein, that interact with FLO2 (Table 1, Figures 3 and ​4);8); however, our results indicate that neither protein interacts with the TPR motif region of FLO2, suggesting that additional interactors may remain to be identified. The LEA proteins, constituting a large gene family in rice, and bHLH proteins are involved in many functions, including a multiple stress response through transcriptional regulation, although their individual functions remain unclear. The flo2 mutant showed reduced expression of multiple genes involved in storage starch and storage proteins (Figures 1 and 2). Therefore, it is suggested that FLO2 might interact with the bHLH protein in a transcriptional complex that could regulate storage starch and storage proteins directly or indirectly through influencing the development of endosperm in rice.

(from reference) .

Figure 3. Phylogenetic Tree of the Representative FLO2 Homologs.(from reference) .

Figure 4. Phylogenetic Tree of the Representative FLO2 Homologs.(from reference) .

In the flo2 mutant, we detected little alteration in the expression of several genes participating in storage starch and storage protein biosynthesis, such as BEIIa, GBSSII, and GluB4 (Figure 1). This result indicates that several genes involved in production of the storage substance may not be regulated by FLO2. Expression profiles of genes involved in starch biosynthesis have been analyzed. Expression patterns of the genes participating in accumulation of storage starch differ depending on the individual genes; GBSSI, SSI, SSIIa, SSIIIa, SSIVb, BEI, BEIIb, AGPL2, and AGPS2b genes are specifically expressed in the endosperm of developing rice seeds at 10 to 20 DAF, whereas GBSSII, SSIIb, SSIIc, SSIVa, BEIIa, AGPL1, and AGPS1 genes are expressed in leaves as well as developing seeds. We observed that expression of all genes in the former group was significantly reduced in the flo2 mutants, whereas little decrease in expression levels was shown in most genes of the latter group (Figure 1). It has been reported that the patterns of gene expression in seeds are tissue and developmental stage specific. Among the genes working in the endosperm, GBSSI, SSIIa, SSIIIa, BEI, BEIIb, AGPS2b, AGPL2, ISA1, and PUL genes, which have transcripts that are low at onset and then rise steeply at the start of starch synthesis in the endosperm, are thought to play essential roles in endosperm starch synthesis at the middle stage of seed development. These genes corresponded to the genes specifically expressed in the endosperm of developing seeds reported by Hirose et al. (2006). Our observation revealed that FLO2 expression was highly elevated at 15 to 25 DAF of seed development (Figure 5). Consequently, it is suggested that FLO2 selectively affects the expression of these genes, whose expression patterns were consistent with that of the FLO2 gene.

Figure 5. Phylogenetic Tree of the Representative FLO2 Homologs.(from reference) .

It has been reported that high-temperature stress during seed development apparently reduces the accumulation of storage starch in rice grains. In the case of Nipponbare, which is a rice cultivar sensitive to high-temperature stress, high temperature during seed development resulted in significant reduction of FLO2 expression, while FLO2 was expressed most abundantly under normal ripening conditions. The timing of elevation of FLO2 expression seemed to be delayed for several days in response to high-temperature stress (Figure 5B). This result suggests that FLO2 expression was apparently repressed by high-temperature stress in this cultivar. On the other hand, in Kinmaze, which is relatively tolerant to high-temperature stress, it seemed that FLO2 expression was upregulated by high-temperature stress (Figure 5A). These facts support the idea that FLO2 plays an important role in the trait for tolerance to high-temperature stress.

FLO2 was abundantly expressed in immature seeds (Figures 6 and ​and 5 ), and its expression pattern was similar to that of genes involved in the production of storage starch and storage proteins. This temporal expression may account for the appearance of the loss-of-function phenotype observed in the endosperm of the flo2 mutant seeds, such as reduced production of these storage substances. While the flo2 mutant produced smaller grains, the OX lines had larger seeds than those of the wild type (Figures 7 and ​8 ), exhibiting the gain-of-function phenotype of the FLO2 gene. The increase in gene expression for enzymes involved in starch biosynthesis, however, is not directly linked to the level of FLO2 expression. As a matter of fact, the increases in some of the parameters of grain size measured are barely significant (Figure 8). This could be attributed to the fact that expression of the analyzed genes was measured at a single time point and that the impact of FLO2 may not coincide with its expression level at that time. This work suggests that FLO2 has a pivotal role in the accumulation of storage substances in the endosperm and sets the foundation for the understanding of grain size and quality regulation in rice.

Figure 6. Phylogenetic Tree of the Representative FLO2 Homologs.(from reference) .

(A) Expression of FLO2 in the stem (S), leaf blade (LB), flag leaf (FL), panicles before heading (P), 10 DAF developing seed (DS), and root (R). Amounts of transcripts are shown as the relative values to those of Actin I. Error bar shows sd (n = 3).

(B) and (C) Expression level of FLL1 (B) and FLL2 (C) in the stem (S), leaf blade (LB), flag leaf (FL), panicles before heading (P), 10 DAF developing seed (DS), and root (R). Closed box, wild type (Kinmaze); open box, flo2 mutant (EM37). Amounts of transcripts are shown as the relative values to those of Actin I. Error bar shows sd (n = 3).

Figure 7. Phylogenetic Tree of the Representative FLO2 Homologs.(from reference) .

Figure 8. Phylogenetic Tree of the Representative FLO2 Homologs.(from reference) .

Evolution

The FLO2 gene belongs to a gene family that consists of FLO2, FLL1, and FLL2 in the rice genome. Orthologs of these genes widely exist in plant species, including mosses, but none was found in yeast or animals. The phylogenetic tree indicates that there are three major clades for higher plants, each of which consists of the orthologs of FLO2, FLL1, and FLL2 (Figure 6). The interspecies similarity between members is larger than the similarities among FLO2, FLL1, and FLL2 within a species. These facts suggest that this gene family was acquired in an ancestor plant and diverged within the plant kingdom. The FLO2 homologs in moss (P. patens) form an independent branch in the phylogenetic tree, suggesting that the moss FLO2 family members may have evolved independently.

Figure 9. Phylogenetic Tree of the Representative FLO2 Homologs.(from reference) .

FLL1 and FLL2 have similar protein structures to FLO2, and presumably have similar functions, but have somewhat different expression patterns. FLO2 was expressed in organs such as leaves and developing seeds (Figure 6A) and is therefore assumed to function in these organs. However, the flo2 mutant showed no apparent phenotype other than in seeds. In both the wild type and the flo2 mutant, a normal level of expression of both FLL1 and FLL2 was observed in leaves, whereas they were poorly expressed in developing seeds. It is presumed that the function of FLO2 is specific to developing seeds.

Labs working on this gene

• Department of Biological Science and Technology, Tokyo University of Science, Noda, Japan
• National Agricultural Research Center, Joetsu 943-0193, Japan
• Institute for Chemical Research, Kyoto University, Uji 611-0011, Japan
• National Institute of Agrobiological Sciences, Tsukuba 305-8602, Japan
• STAFF Institute, Tsukuba 305-0854, Japan
• Institute of Genetic Resources, Faculty of Agriculture, Kyushu University, Fukuoka 812-8581, Japan

Structured Information

Gene Name	Os04g0645100
Description	Tetratricopeptide-like helical domain containing protein
Version	NM_001060598.1 GI:115460925 GeneID:4337192
Length	11244 bp
Definition	Oryza sativa Japonica Group Os04g0645100, 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 4
Location	Chromosome 4:33233812..33245055
Sequence Coding Region	33233846..33233930,33234571..33234632,33234925..33235017,33235313..33235385,33238210..33238515 ,33238601..33238659,33238772..33238903,33239038..33239079,33239184..33239717 ,33239813..33240003,33240162..33240651,33240897..33240959,33241328..33241429 ,33241519..33241839,33241918..33242010,33242108..33242209,33242342..33242446 ,33242557..33242690,33243053..33243236,33243850..33243981,33244071..33244183 ,33244252..33244333,33244952..33245055
Expression	GEO Profiles:Os04g0645100
Genome Context	<gbrowseImage1> name=NC_008397:33233812..33245055 source=RiceChromosome04 preset=GeneLocation </gbrowseImage1>
Gene Structure	<gbrowseImage2> name=NC_008397:33233812..33245055 source=RiceChromosome04 preset=GeneLocation </gbrowseImage2>
Coding Sequence	<cdnaseq>atggcccccaagggcgccggccgtggcaagggcaggggcggcggcggcggcggcaaaggcgacaagaggaagaaggaagagaaagttgtgccgagtgccatagacgtcacggttgtcactccctatgaatcccaggttacactcaaggggatatctacggaccgtgttctggacgtgaggaagctgctgggctcgaatgtggagacgtgccacctgacaaactactcgctctcgcacgtgacgcgagggcagcggctggaggacggcgtggaaatcgtgtcgctgaaaccatgctctctcaccatcgtggaagaggagtacgcgacggaggcggcggcggcggcgcaagtccggcggctgctggacatcgtcgcctgcaccaccgcgttcgtgaacaagccccgcgacggcgccaagcacaagtcgtccaagcacgcgcgcccggcgacgccgccgtcaccgcccgcgctcgccgcctcccccgacgcccatggcgccggcgccgcccaggcgccgccgatatcggaggcgcacgacatggcggccatccgtccgccgcccaggctgggcgagttctacgacttcctctccttcgcccacctcaccccgcccgtccactttatcaggaggaaggagagcaatggcgcttctcaggaaggcgactacttcgaaatcgaggtgaaagtttgcaatgggaagctcctgcacatcgttgcttccgtcaagggcttctattcggcagggaagccgcacaccgtgagccactccttggtggatctgttgcaacagctcagcagtgcatttgccaatgcatatgatgcactgatgaaagcttttctggatcacaacaagtttgggaatctaccatacggatttcgtgcaaacacatggcttattcctcctatatacttggattctgctacaaagtgcccagcattaccagttgaggatgagaattggggtggaaacgggggtggcaatgggcgagatggaaagtatgacagaagacgatgggcaaaagaattttctactttggctagaatgccatgcaaaactgaggaagggagggtgatcagagaccggaaggcttttcttttacacaatctctttgtagatacagcaatttttagagctgcatcaacaatacagagactcattgatctgtctgggaactcaactagtcaacaagctggcctagatggttctttagcaatcgaggaacgtgttggtgacttgcttataactgtaaagagggatcaggctgatgctagtttaaagctggaggataaagttgatggagttgcactctatcaaaccggttctatggatatatcacaaagaaatcttctaaaaggtttaacttctgatgaaagtgtcgttgttaaggacacctcaatactcggagtggttattgttaaacattgtggatatacagcaacagtgaaggtttcagggcgcacaaaggacggaaatggtggcaaacaaaccagtgatatctgtgatcatctcgatggaatttcgaatgttgatgtagatgatttgccagatggaggttctaatgccttgaacattaacagtctaagaatatcactgccaaaaattgtcaactcagacatcgctagtactcagtgtcctactccgcagtctcatgttgataatcatgcaaggaaactagtgcgcaaaatacttgaagatagcttgatgaagctggagaacatgcctgccaataatcctagaaccatcagatgggaacttgggtcatcctggctgcaaaacttgcagaaaaaggattcgccggcatctgaggacaagaaaaatgcaggacatgttgaaaaagagacaactatcaaaggtcttggtaagcactttgaacagctaaagaaaattaaaaagaaagaatgccatgttgaaggtgctatgtctgagaaggaagactctgacagtaactgctctgtcataaatggcatggaagaatctgaaaacaccaaggagactgatattagcaagctgatgtcagaggatgatttctgtcgccttaaggatctgggagctggacttcatcaaaagtcccttgaagagctcactatgatggcccataaattctatgatgatactgcgcttccaaagctggtggctgattttgcttcccttgaactttctcctgtggatggaagaaccatgaccgatttcatgcacacaaggggtcttaatatgtgctccttaggccgtgtggtagagctcgcggagaagcttccacatatccagtcaatttgcatccatgaaatggtcattaggtcctttaagcacattgttcgagctgttattgcagctgttgatgacatgcaaaatatgtctgcagctattgctgagactttaaacattttacttggatgcccaagacttgaaagtggcactgaaacagatgcacatagtgaacataatttgagatttagatgggtagagaggttcctttctaaaaggtacaattggaaattgaaagatgaatttgcacacttgcggaaatttatcattctgagaggtctttgcagtaaggttggacttgagttggttgcgagagattatgatatgaatagcccaaatccatttgacaaatctgacatagttaacattattcctgtatgcaagcatgtggtgtactcttctattgatggtcgaaacttattagaatcatcaaagatggctttggataaaggaaaacttgatgatgccgtcaacttcggaacaaaggcgttgtccaaaattgtagcagtttgtggtccataccatcggctgactgctaatgcgtacagtcttcttgctgtagttctttaccatactggagattttaatcaggcaactatatatcagcagaaggcacttgatatcaatgaaagagagctaggtcttgaccacccagaaactatgaagagttacggggatttatctgtcttctactaccgtctccagcacatcgaaatggctctaaagtatgtcaaccgtgcactatatctactccaattttcttgtgggctttcacatccaaattcagctgccacatacataaatgtggctatgatggaagaaggtatgggaaatgttcatgtcgctctcaggtacttgcatgaagcactgaagtgcaacaagagactgctaggagctgatcacattcagactgctgcgagctatcatgccatagccatagccctctcaatgatggatgcatactccttgagtgttcagcatgaacagaccaccttacagatacttcaggagaaactaggacaagatgaccttcgaactcaggatgctgctgcatggcttgagtattttgagtcaaaagcattagaacaacaagaggccgcccgaaggggcatcccaaaacctgactcatctattgcaagcaaaggacatcttagtgtatcagatctacttgactacataagcccggatcaggaaagaaaagagagggatacacaaagaaagggcaggcgtgcaaagaataatatcagagcccatcaaggtgaattagttgaagaaaaggagagctttgagcacgacataggatctcctcatgaagcaaacaaagaggagtttcaacaagt</cdnaseq>
Protein Sequence	<aaseq>MAPKGAGRGKGRGGGGGGKGDKRKKEEKVVPSAIDVTVVTPYES QVTLKGISTDRVLDVRKLLGSNVETCHLTNYSLSHVTRGQRLEDGVEIVSLKPCSLTI VEEEYATEAAAAAQVRRLLDIVACTTAFVNKPRDGAKHKSSKHARPATPPSPPALAAS PDAHGAGAAQAPPISEAHDMAAIRPPPRLGEFYDFLSFAHLTPPVHFIRRKESNGASQ EGDYFEIEVKVCNGKLLHIVASVKGFYSAGKPHTVSHSLVDLLQQLSSAFANAYDALM KAFLDHNKFGNLPYGFRANTWLIPPIYLDSATKCPALPVEDENWGGNGGGNGRDGKYD RRRWAKEFSTLARMPCKTEEGRVIRDRKAFLLHNLFVDTAIFRAASTIQRLIDLSGNS TSQQAGLDGSLAIEERVGDLLITVKRDQADASLKLEDKVDGVALYQTGSMDISQRNLL KGLTSDESVVVKDTSILGVVIVKHCGYTATVKVSGRTKDGNGGKQTSDICDHLDGISN VDVDDLPDGGSNALNINSLRISLPKIVNSDIASTQCPTPQSHVDNHARKLVRKILEDS LMKLENMPANNPRTIRWELGSSWLQNLQKKDSPASEDKKNAGHVEKETTIKGLGKHFE QLKKIKKKECHVEGAMSEKEDSDSNCSVINGMEESENTKETDISKLMSEDDFCRLKDL GAGLHQKSLEELTMMAHKFYDDTALPKLVADFASLELSPVDGRTMTDFMHTRGLNMCS LGRVVELAEKLPHIQSICIHEMVIRSFKHIVRAVIAAVDDMQNMSAAIAETLNILLGC PRLESGTETDAHSEHNLRFRWVERFLSKRYNWKLKDEFAHLRKFIILRGLCSKVGLEL VARDYDMNSPNPFDKSDIVNIIPVCKHVVYSSIDGRNLLESSKMALDKGKLDDAVNFG TKALSKIVAVCGPYHRLTANAYSLLAVVLYHTGDFNQATIYQQKALDINERELGLDHP ETMKSYGDLSVFYYRLQHIEMALKYVNRALYLLQFSCGLSHPNSAATYINVAMMEEGM GNVHVALRYLHEALKCNKRLLGADHIQTAASYHAIAIALSMMDAYSLSVQHEQTTLQI LQEKLGQDDLRTQDAAAWLEYFESKALEQQEAARRGIPKPDSSIASKGHLSVSDLLDY ISPDQERKERDTQRKGRRAKNNIRAHQGELVEEKESFEHDIGSPHEANKEEFQQV</aaseq>
Gene Sequence	<dnaseqindica>35..119#760..821#1114..1206#1502..1574#4399..4704#4790..4848#4961..5092#5227..5268#5373..5906#6002..6192#6351..6840#7086..7148#7517..7618#7708..8028#8107..8199#8297..8398#8531..8635#8746..8879#9242..9425#10039..10170#10260..10372#10441..10522#11141..11244#atttgtggctgttgggtttggagcttcaggcgcaatggcccccaagggcgccggccgtggcaagggcaggggcggcggcggcggcggcaaaggcgacaagaggaagaaggaagagaaaggtattgtgtagtacatgttgctggaagctcgaaaccgcttgtgctctgcaaaggcatagagttcacttcatggcatgaagctgtgttgctgttgccgtgtgaggagctttgttcttgcgaacattgaattcgctgctgctgctgctgctgctgtaatctgtgtgtgtttgctgttctggcttgggttcttttgcatgggtaaagggtgaaaaaaagacgatggtctctctgctattagtagcggtaatcttttctttattccttcttttgcttttgtttctcttccttttttcttgcctattcgtactcctgatttcggtgattatattgctgtttaagcgcttttttcttggtgtttagcttcgtgttctgtgctcatgattattactactagtcttttgtcttttttttttttgcaccttctcttgttccattaatccttttcgttcgcttccaagcaacatggcccgattctttccttggtgtaacgcacaccgtcgtgcggttgagtcatgcgttctgcgagctctagatgaatactctgttttcactttcacctcgacatgatgtcgccgtctccgtttttaaggtttttttctgcagctgttggtatgatgctgggttcttcttgtttttgcagttgtgccgagtgccatagacgtcacggttgtcactccctatgaatcccaggttacactcaaggtatgtgtggctgcattagctggctggtacttgctgcacacatgaacacacaaaatacctgaaaggacaatggacaggacaagcaaaagtgaacagaaactggtatctttgggaaaagggggcctaattctgataactgaaatgtctgcagctatttgcaatgagctgcaaagccagttattacttactccacactgcatgctcacaatgagtccgggaaattgagcctttgaaatggaacagtgaacctgagtgaaaagattaatcttgtttttgttctgtgtgtgttcaggggatatctacggaccgtgttctggacgtgaggaagctgctgggctcgaatgtggagacgtgccacctgacaaactactcgctctcgcacgtggtaatgcttgcttcttgtctgctaattcaatcaattgcagtgcttgcagtatctactgataattcaattgcagtgctagtactagcatctatcattccactcttaagtcaaagtgttaattactacttaattatctgccacttctttgctactcttctggaattcacccgggattaatttgaggacagtaacttttaatgtatgtgtgctgcaatgctgttaattctgtaactaattctgtataagctgtgttcggtgaagtctgaatttgtgaagcgtgtgatgtatgacgcagacgcgagggcagcggctggaggacggcgtggaaatcgtgtcgctgaaaccatgctctctcaccatcgtggaaggtacggtccatggtgaaagaatccgcacgctagctgctcttcacttgttcttccttgctcgttttgtttctttctcctttctttttttttcgcttggcaacatggaatgcttcacataacatgatcgaggtggggtagctggcattaggacacataattagttggatgcttaagtagacgaagcaacctgacgagaggacaaaattaagcttgagcttgtggtgtaccgtgtggcacaatctgttagcatgacagtgttatgctacagtccaagtgtccaaacagtctgccatcaatgattaatcaaccaacggtccggatttttccactgaaagctctaaagtctgacgctaatattgctaatattagggaagaacctaatatcaaataattagaaggggtgaggcttcaaatctaggtcgtctagtccaccatcttgtggagctagctggaagacccctaagtgtttctcataagctctaaagtctgaacatgcgtttgccgttgtccgatggtgaagctagtggtttttgttaatgttgactcaccggtagatcatatcaaacctaacttcaaaaatcataatctattcaaattgggacggcacaatgcacatagatgcagccttgtttgtcgacgcacagcattgaatcgaagagcatggaccgcgccggcctttcggctgaataaacttcagaagaaaaaaacacacacatatgactagatcactagattcgcatgctaaggtgcaacctcccacgactccttgtaaatgcagtcatgcgcctccatcacgccccaatactggcacaccagtgagttagataggagcacagagtacaatctagttcattatacttcataattcttgccacttagacagcgagacgtgcaggcaggcagcagcagcagctttgcatagtgcctccagtttcagacattagaagcagcgcatcctgtttcagacacttagcataaaattgtttagaccaaaaaaaatgtgatttacatctcgggtctctaccgttatcctcagatactagcagcaaatggtagccggtttcagtgccgcatctgtagccgatgggagatatagatcgtggccgtgacgtcgtcgcgcgatctgcctctccgcggttggttggggcggggcggctaaggtgttttttccacacacacacacatgtcccgtctctttcgctgccgatgtcgcatttgccgtcgccgcgcgaacgtttcacggcggctttgccagtttccgggaggaaggaggcggcggttgaaccgaacctggacgcccccgcgcgcacgccggctaataagctgaggccctgtttagatgggactaaaacacttttaaaaacacttttaagttcctatcacatcggatgtgtactaattataaatattaaatgtaactattaataaaatctatccataatcttggactaattcgcgagacgaatctatttgagcctaattaatccatgattagcctatgtgatgctacagtaaatattctctaattatggattaattaggcttaaaaaatttgtctcgcgaattagctttcatttatgtaattagttttgtaagtagtctatatttaatactctaaattagtgtctaaagatcagatatggactaaagttaagtccctggatccaaacactcgctgacggatgccggtgatgtttcgtatggttttttttcaacgaaacgtgtggcagcgatggtcgtggcatgcgaccatgtgtctgtgtcgacgacgaaaacgagcatagaacttaacgattgattagtgcgtaagactgtacgaggaaggggatgactgaccgattggtctgacacgcgctcgtagtcaaatgcacttgcttttactgctaccgtagtccatcattttaaaaaaaaaaacctgaaattcaagtgcaaagtggcatttggacaaactggaagggcattttcactcgttgatagcgcttgattagttgcctaggcgaagtccgtcttgtgcacttctctagtgggtcagagcctcttgttctgctttgggaaatacatgctcgttaattaactgaacatagtgctccacctgctgtctgtctcatatggtatttagaactgaaagctatactagtgtacaattaattactgcagaattatgcgattttgtagttgttgtatgctccaaggtctgagctaatggctcaccggttgttgtcctgtcctaaacgccggtacgggacaaaaggacgcctcttggaatcagcgttgggcgttagctcaagtgcctaacttattttactcgccactcttctgacgaatcgacgggtttgtgaattgtgattgtacatttgtttgtagtattggtgtttaggtggtcactgactcatactggtcagtacagtacgtgctagacctgctaccggtcgtaccgatgccatgagtcaaaatcacgttgttggccatgtgtttctaactcgctgacaccgctgggaacgtgatagaatagcgacatagcggtgatagtacttactactatgacacgagtatcaaaacttcagccacgttcctgtggatgaacatcagtcttaagccattttttttttatttttacatttgaattcgatttattctattttgatttgattcgacgacttttgttcaatatgatgcagcagctgctaaaaaaacaaaagaaattgcttcgcttatttgttccagaattgatgggcctgttcatttataaagggaagcccagcacttggcctattgggcctgaattttggtagagctaatcgagcccatgggcctttttgtgcagaggagtacgcgacggaggcggcggcggcggcgcaagtccggcggctgctggacatcgtcgcctgcaccaccgcgttcgtgaacaagccccgcgacggcgccaagcacaagtcgtccaagcacgcgcgcccggcgacgccgccgtcaccgcccgcgctcgccgcctcccccgacgcccatggcgccggcgccgcccaggcgccgccgatatcggaggcgcacgacatggcggccatccgtccgccgcccaggctgggcgagttctacgacttcctctccttcgcccacctcaccccgcccgtccactgtaaggccgaatgctgcatcgcgtttccccctttttgcgaggcgtggaactaggggttggactgacgcgtggttgcggtgtgcagttatcaggaggaaggagagcaatggcgcttctcaggaaggcgactacttcgaaatcgaggtgtgatgcgatgtctctcagaacgctagagagttcgtctgatctgcagtagccaactccgcgatgagtctatgactacgcacttcactgacgggagttgatgtgtggccaggtgaaagtttgcaatgggaagctcctgcacatcgttgcttccgtcaagggcttctattcggcagggaagccgcacaccgtgagccactccttggtggatctgttgcaacagctcagcagtgcatttgccaatgtgagtaaccatgaatcctgaacatgtataatgtataacgcgttgctcgatttactttctgctcctgtttgtttttggctaactatacagccgaccttatctgacagtgatgtcttctaattcattttgcacaggcatatgatgcactgatgaaagcttttctggatcacaacaaggtggttatagtaaatgctgtttcctttttttttttcccccttataccgtagatttagtcaagtaaaatgctaaatttgacaaggatacttctttgcatctccagtttgggaatctaccatacggatttcgtgcaaacacatggcttattcctcctatatacttggattctgctacaaagtgcccagcattaccagttgaggatgagaattggggtggaaacgggggtggcaatgggcgagatggaaagtatgacagaagacgatgggcaaaagaattttctactttggctagaatgccatgcaaaactgaggaagggagggtgatcagagaccggaaggcttttcttttacacaatctctttgtagatacagcaatttttagagctgcatcaacaatacagagactcattgatctgtctgggaactcaactagtcaacaagctggcctagatggttctttagcaatcgaggaacgtgttggtgacttgcttataactgtaaagagggatcaggctgatgctagtttaaagctggaggataaagttgatggagttgcactctatcaaaccggttctatggatatatcacaaagaaatcttctaaaaggtttaacttctgatgaaagtgtcgttgttaaggtggtggcatactactttcatatctctttttttttcaaattattttttgtccttaatattaattaccgatttccaccaacatgattaccattcaggacacctcaatactcggagtggttattgttaaacattgtggatatacagcaacagtgaaggtttcagggcgcacaaaggacggaaatggtggcaaacaaaccagtgatatctgtgatcatctcgatggaatttcgaatgttgatgtagatgatttgccagatggaggttctaatgccttgaacattaacaggtttgatatatgttcccatgtaaacttattcacatgcttgctcatcagtagagaaaataatgttcactggtaactagtttttcttacatttggcacacaataatgttccattaacacaactctctctctctctctctctctgtatatgtgtgtttcagtctaagaatatcactgccaaaaattgtcaactcagacatcgctagtactcagtgtcctactccgcagtctcatgttgataatcatgcaaggaaactagtgcgcaaaatacttgaagatagcttgatgaagctggagaacatgcctgccaataatcctagaaccatcagatgggaacttgggtcatcctggctgcaaaacttgcagaaaaaggattcgccggcatctgaggacaagaaaaatgcaggacatgttgaaaaagagacaactatcaaaggtcttggtaagcactttgaacagctaaagaaaattaaaaagaaagaatgccatgttgaaggtgctatgtctgagaaggaagactctgacagtaactgctctgtcataaatggcatggaagaatctgaaaacaccaaggagactgatattagcaagctgatgtcagaggatgatttctgtcgccttaaggatctgggagctggacttcatcaaaaggtaatttatagctatacagttcatcaatcaatatggtctgaacaatttagttttatgtccggtaagctcatacatgctattttgcttgggattaaagttctaatctgttaaaattttcaccagaagccttgtgttaagctgaatatgatgtagtagcagaagcgcagcattgtatttttccagtaatcaggaatatattggttattttgctgtttaatacttaactttgttatttcttgaactagtcccttgaagagctcactatgatggcccataaattctatgatgatactgcgcttccaaagctggtaagccttgccttgtttatttggagatccttatgtaaattctcgaacattttaatcggcactctcttctctgaaaacaaagtagtcgctttttgctggcattggtattgtttattgtttattgttgtagtaagtagtttagttatggtactaatgttcagttagttcatttgtaattttgaggaagtagtcaatccacattaacaattcgcaccacaaagtccaaaagaattacagttcttgaaaatttttgcgtgtaaagaagaaaaaaaaaacaatgttttttttatgttgtcttaaactttcttaattattatttgatgcccataatagatagttaattttttcttggaataaaataatgacaggtggctgattttgcttcccttgaactttctcctgtggatggaagaaccatgaccgatttcatgcacacaaggggtcttaatatgtgctccttaggccgtgtggtgagtttaaatttattctgaattacttttgcttctatggttggatggaatttccagaataaccttttttttatatataaattaaacaggtagagctcgcggagaagcttccacatatccagtcaatttgcatccatgaaatggtcattaggtcctttaagcacattgttcgagctgttattgcagctgttgatgacatgcaaaatatgtctgcagctattgctgagactttaaacattttacttggatgcccaagacttgaaagtggcactgaaacagatgcacatagtgaacataatttgagatttagatgggtagagaggttcctttctaaaaggtacaattggaaattgaaagatgaatttgcacacttgcggaaatttatcattctgagaggtctttgcagtaaggtacaatccaaagtgtgttccaaaacattcatagagtacagtttctgaacatgcataattaatatgtttatttctcaggttggacttgagttggttgcgagagattatgatatgaatagcccaaatccatttgacaaatctgacatagttaacattattcctgtatgcaaggtaggacactatcagaaaacttaccactgtactttccccaccacctgaattaataattatattgatttgatctgctttggccttgcgcttcttacagcatgtggtgtactcttctattgatggtcgaaacttattagaatcatcaaagatggctttggataaaggaaaacttgatgatgccgtcaacttcggaacaaaggtatgcaataatcagttcttttttctgtacgttgtttaacagctatgctaaaatatttctcttcctgctttggttgggtgaagttaccattctaaattctttcattaactatttttttttcttttcttgaaggcgttgtccaaaattgtagcagtttgtggtccataccatcggctgactgctaatgcgtacagtcttcttgctgtagttctttaccatactggagattttaatcaggtatttcaatgaatttggtgggctgtattctatctgctgcatagtgatgtgtttttggcacaatgtttggcaactgcatttttcactagacttctgttctttctctgtaggcaactatatatcagcagaaggcacttgatatcaatgaaagagagctaggtcttgaccacccagaaactatgaagagttacggggatttatctgtcttctactaccgtctccagcacatcgaaatggctctaaagtaagtgctgctattaggtgttttgttgtggaaacacaaaagctagtttcatgacttggcttgtagctggtattactatcatgcacaatgtcgttacgctggttcatttctagccattacatgcattaaatatattgcctgtagaatggatactactgcatttgtacggcactatggctgtattaatttgtagacttgaactgtgcattgttttgcattattgtccatgattgttacttgtggtttgcacggaaacttgctgaaggcacatatttgcttgacaatattgtcaaaactcatcttccatctttctttttttgccttttgttttttgtatcgctaatggtatctttttcttaaaggtatgtcaaccgtgcactatatctactccaattttcttgtgggctttcacatccaaattcagctgccacatacataaatgtggctatgatggaagaaggtatgggaaatgttcatgtcgctctcaggtacttgcatgaagcactgaagtgcaacaagagactgctaggagctgatcacattcaggtaaacaaaactacatatttttctcttaccgtgctttcacagctttatctgttttcctggaaacaataatctaattgttttttttgttggacttgattgatgtgcattaaagtacagtcttgtaagatcaattccatgcgtactatctcttgcaccaatatttctgcagaagtactggcattggcggcatacttgtcttattaaggatattagacatatacacctctgatacagtagaaatgtcccataagtatctatgtctgaagcaaactttttgatttagctctaatatccctagatattctccattgctgtccctcaaaggccatgagcaattgtttaagtttctttcaatttcctatccaacttgtttattgcatatccaatttcaaaactagcatcttgtttcagtagtacccgcgccaccatgacttgtatcttttaaggaaaggtttattttgatgtctagacccacagaaacagtcttttctgtatatttcaatccgtttgacataatgcatgtaactacggacaacttcaatggcgtatttgttatgcatatgatcccgtgattcagttaacatgaattttcatgcatttctttgaacaacagactgctgcgagctatcatgccatagccatagccctctcaatgatggatgcatactccttgagtgttcagcatgaacagaccaccttacagatacttcaggagaaactaggacaagatgaccttcgaactcaggttttttttcccttcaaaatatgcaatgacgagtccaaatactgctatgttctcttggactattagacatgataattttcctgctgcaggatgctgctgcatggcttgagtattttgagtcaaaagcattagaacaacaagaggccgcccgaaggggcatcccaaaacctgactcatctattgcaagcaaaggacatcttaggtatgaatacactgtgcattatcagtcgcaaactgtgatgatctggattgaaacatttatctttccagtgtatcagatctacttgactacataagcccggatcaggaaagaaaagagagggatacacaaagaaagggcaggcgtgcaaaggtgattataacatgcctactttttcaaaattattgttgttaaaatgtagtcagaagagataagtgcacactggacagtggagacaattccatcagtattcatgaaaagtctagatgagaataggattggcacaataacttccatataggatgtctggtatatgaactgatgaacatatgctggcagcaaatcagcaatcctatccagttcatgaaactagtaaaaggttatgaaccatcatcactgacataataaccatgacagttatccccaccagtcattgttggagcatgaactatgagaattgaaaagatattcccgcatccacagctagtccattgataccaggcatttatccagatgattccagttcaacccattaacttagcaatgaattgacaacacaatttcttcttagttgggaatttaactccctttacatatggatatttaatcctttgttttctaatatgcttcatgaattgggaatcacaatatacatattttggaagtccattgatgtcatggtttcaattgttgcctaaacattatgcaagcttttattttacccttactaccctttgctttccttttcaccttgtatagtattaaatgcagaataatatcagagcccatcaaggtgaattagttgaagaaaaggagagctttgagcacgacataggatctcctcatgaagcaaacaaagaggagtttcaacaagt</dnaseqindica>
External Link(s)	NCBI Gene:Os04g0645100, RefSeq:Os04g0645100