Difference between revisions of "Os07g0211500"

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    Rc is a domestication-related gene required for red pericarp in rice (Oryza sativa) and it encodes a basic helix-loop-helix (bHLH) protein that was fine-mapped to an 18.5-kb region on rice chromosome 7 using a cross between Oryza rufipogon (red pericarp) and O. sativa cv Jefferson (white pericarp).
+
Rc is a domestication-related gene required for red pericarp in rice (Oryza sativa) and it encodes a basic helix-loop-helix (bHLH) protein that was fine-mapped to an 18.5-kb region on rice chromosome 7 using a cross between Oryza rufipogon (red pericarp) and O. sativa cv Jefferson (white pericarp).
 
===characteristic===
 
===characteristic===
    Different colors, such as purple, brown, red and white, occur in the pericarp of rice. Here, two genes affecting proanthocyanidin synthesis in red- and brown-colored rice were elucidated, one of the gene is Rc.<ref name="ref5" />)
+
Different colors, such as purple, brown, red and white, occur in the pericarp of rice. Here, two genes affecting proanthocyanidin synthesis in red- and brown-colored rice were elucidated, one of the gene is Rc.<ref name="ref5" />)
 
===Mutation===
 
===Mutation===
 
[[File:Clip image002.gif|right|thumb|150px|''picture1. Length mutation (from reference <ref name="ref1" />).'']]
 
[[File:Clip image002.gif|right|thumb|150px|''picture1. Length mutation (from reference <ref name="ref1" />).'']]
[[File:Clip_image0021.gif]]
+
[[File:Clip_image0021.gif]]Grain color has been the target of selection in domesticated Asian rice (as well as other domesticated grains), and the vast majority of O. sativa varieties have white (nonpigmented) grains, compared to the dark red grains that characterize the wild relative and other wild species. The genetic basis of white grains in O. sativa has been pinpointed to loss-of-function mutations in the Rc gene, a regulatory protein in the proanthocyanidin synthesis pathway; these loss-of-function mutations prevent the development of a pigmented pericarp layer. A survey of 337 white pericarp rice cultivars showed that the majority (>97%) have a 14-bp deletion in exon 7 of the gene, resulting in a premature stop codon and a non-functional allele referred to as rc.
    Grain color has been the target of selection in domesticated Asian rice (as well as other domesticated grains), and the vast majority of O. sativa varieties have white (nonpigmented) grains, compared to the dark red grains that characterize the wild relative and other wild species. The genetic basis of white grains in O. sativa has been pinpointed to loss-of-function mutations in the Rc gene, a regulatory protein in the proanthocyanidin synthesis pathway; these loss-of-function mutations prevent the development of a pigmented pericarp layer. A survey of 337 white pericarp rice cultivars showed that the majority (>97%) have a 14-bp deletion in exon 7 of the gene, resulting in a premature stop codon and a non-functional allele referred to as rc.
+
Our findings for the pericarp color gene, Rc, indicate that remarkably similar genetic mechanisms can be at play. We find that two of the three white pericarp O. glaberrima varieties in our sample harbor a unique point mutation that is predicted to result in a premature stop codon in exon 7 of the Rc gene.
    Our findings for the pericarp color gene, Rc, indicate that remarkably similar genetic mechanisms can be at play. We find that two of the three white pericarp O. glaberrima varieties in our sample harbor a unique point mutation that is predicted to result in a premature stop codon in exon 7 of the Rc gene.
 
 
Instead, two samples contained a novel point mutation predicted to result in a premature stop-codon in exon 7, hereafter referred to as allele rc-g1 . This mutation is 146 bp upstream of the Rc-s point mutation and 201 bp upstream of the initiation of the 14-bp rc deletion. The third white pericarp O. glaberrima contained no identifiable insertions, deletions, or premature stop-codons, nor does it differ from the red-pericarp O. glaberrima sequences for any predicted amino-acid changes. Comparison of this sample to the most closely related red-pericarp sample showed that they were identical for the sequenced region upstream of the start codon (>1.5 kb) and for the first intron; both of these regions are potentially important as cis-regulatory regions in plants.
 
Instead, two samples contained a novel point mutation predicted to result in a premature stop-codon in exon 7, hereafter referred to as allele rc-g1 . This mutation is 146 bp upstream of the Rc-s point mutation and 201 bp upstream of the initiation of the 14-bp rc deletion. The third white pericarp O. glaberrima contained no identifiable insertions, deletions, or premature stop-codons, nor does it differ from the red-pericarp O. glaberrima sequences for any predicted amino-acid changes. Comparison of this sample to the most closely related red-pericarp sample showed that they were identical for the sequenced region upstream of the start codon (>1.5 kb) and for the first intron; both of these regions are potentially important as cis-regulatory regions in plants.
 
This study also shows that white pericarps might be achieved by mutations in cis-regulatory regions of Rc or potentially mutations in other genes, a pattern that was not observed in an extensive survey of O. sativa cultivars.(from reference <ref name="ref1" />)
 
This study also shows that white pericarps might be achieved by mutations in cis-regulatory regions of Rc or potentially mutations in other genes, a pattern that was not observed in an extensive survey of O. sativa cultivars.(from reference <ref name="ref1" />)
    Crop domestication can serve as a model of plant evolutionary processes. It involves a series of selection events from standing natural variation and newly occurring mutations and combinations of mutations as a result of natural crossings in populations during local adaptation and propagation of plant lines to other cultivation areas.<ref name="ref3" />)
+
Crop domestication can serve as a model of plant evolutionary processes. It involves a series of selection events from standing natural variation and newly occurring mutations and combinations of mutations as a result of natural crossings in populations during local adaptation and propagation of plant lines to other cultivation areas.<ref name="ref3" />)
    To determine the subpopulation origin of the Rc mutations,we examined ancestral haplotypes across the Rc coding sequence and promoter region in 103 genetically diverse, redgrained rices. Four rice insertion/deletion polymorphism (RID) and two rice microsatellite (RM) polymorphisms were used to construct haplotypes across the 6.5-kb region containing the Rc gene.<ref name="ref4" />)
+
To determine the subpopulation origin of the Rc mutations,we examined ancestral haplotypes across the Rc coding sequence and promoter region in 103 genetically diverse, redgrained rices. Four rice insertion/deletion polymorphism (RID) and two rice microsatellite (RM) polymorphisms were used to construct haplotypes across the 6.5-kb region containing the Rc gene.<ref name="ref4" />)
  
 
===Function===
 
===Function===
[[File:Struture1.jpg]]
+
[[File:Struture1.jpg]]Rc alleles from the temperate ecotype of weedy lines LD and C9541 could represent the functional alleles most distant to those in group I (Figure 4). All the donor lines of weedy and cultivated rice in both groups were identified as dormant genotypes, or dormancy was associated with the Rc locus in our previous  or present research. Thus, phylogenetic and dormancy data suggest that Rc alleles functional for pericarp color most likely also function for seed dormancy.<ref name="ref2" />)
    Rc alleles from the temperate ecotype of weedy lines LD and C9541 could represent the functional alleles most distant to those in group I (Figure 4). All the donor lines of weedy and cultivated rice in both groups were identified as dormant genotypes, or dormancy was associated with the Rc locus in our previous  or present research. Thus, phylogenetic and dormancy data suggest that Rc alleles functional for pericarp color most likely also function for seed dormancy.<ref name="ref2" />)
 
  
 
===Expression===
 
===Expression===
    RT-PCR experiments confirmed that the Rc gene was expressed in both red- and white-grained rice but that a shortened transcript was present in white varieties.We anticipated that similar expression levels of Rc would be detected in red and white seeds. Our results confirmed this expectation and further demonstrated that the RNA transcript from cv Jefferson contained the 14-bp deletion predicted from the sequence information.(from reference <ref name="ref6" />)
+
RT-PCR experiments confirmed that the Rc gene was expressed in both red- and white-grained rice but that a shortened transcript was present in white varieties.We anticipated that similar expression levels of Rc would be detected in red and white seeds. Our results confirmed this expectation and further demonstrated that the RNA transcript from cv Jefferson contained the 14-bp deletion predicted from the sequence information.(from reference <ref name="ref6" />)
  
  
 
===Nucleotide Polymorphisms===
 
===Nucleotide Polymorphisms===
    Phylogenetic analysis, supported by comparative mapping in rice and maize (Zea mays), showed that Rc, a positive regulator of proanthocyanidin, is orthologous with INTENSIFIER1, a negative regulator of anthocyanin production in maize, and is not in the same clade as rice bHLH anthocyanin regulators.[[File:mapping of Rc.jpg]](from reference <ref name="ref6" />)
+
Phylogenetic analysis, supported by comparative mapping in rice and maize (Zea mays), showed that Rc, a positive regulator of proanthocyanidin, is orthologous with INTENSIFIER1, a negative regulator of anthocyanin production in maize, and is not in the same clade as rice bHLH anthocyanin regulators.[[File:mapping of Rc.jpg]](from reference <ref name="ref6" />)
 
===Allele Distribution===
 
===Allele Distribution===
    Sequencing of the alleles from both mapping parents as well as from two independent genetic stocks of Rc revealed that the dominant red allele differed from the recessive white allele by a 14-bp deletion within exon 6 that knocked out the bHLH domain of the protein.(from reference <ref name="ref6" />)
+
Sequencing of the alleles from both mapping parents as well as from two independent genetic stocks of Rc revealed that the dominant red allele differed from the recessive white allele by a 14-bp deletion within exon 6 that knocked out the bHLH domain of the protein.(from reference <ref name="ref6" />)
  
 
===Recombination===
 
===Recombination===
    The rg7.1 locus was originally mapped to a 7.2-Mb region that included the centromere on rice chromosome 7. Given the low frequency of recombination across this region, it was not clear whether positional cloning would be feasible. This study demonstrates that even in regions that are recombinationally repressed, map-based gene isolation offers a viable approach.(from reference <ref name="ref6" />)
+
The rg7.1 locus was originally mapped to a 7.2-Mb region that included the centromere on rice chromosome 7. Given the low frequency of recombination across this region, it was not clear whether positional cloning would be feasible. This study demonstrates that even in regions that are recombinationally repressed, map-based gene isolation offers a viable approach.(from reference <ref name="ref6" />)
 
===Evolution===
 
===Evolution===
    Repeated phenotypic evolution can occur at the interspecific level, where it is manifest as the appearance of the same trait in multiple domesticated species, and at the intraspecific level, where the same trait arises multiple times within a single crop.
+
Repeated phenotypic evolution can occur at the interspecific level, where it is manifest as the appearance of the same trait in multiple domesticated species, and at the intraspecific level, where the same trait arises multiple times within a single crop.
 
In rice, the opportunity exists to examine the genetic basis of repeated trait evolution at both the interspecific and intraspecific level.
 
In rice, the opportunity exists to examine the genetic basis of repeated trait evolution at both the interspecific and intraspecific level.
 
At the intraspecific level, repeated phenotypic evolution during domestication can be explored in two contexts; one is the repeated evolution of a trait in varieties resulting from a single domestication event, the other is repeated evolution of a trait in varieties that result from multiple, independent domestication events within the same species.(from reference <ref name="ref1" />)
 
At the intraspecific level, repeated phenotypic evolution during domestication can be explored in two contexts; one is the repeated evolution of a trait in varieties resulting from a single domestication event, the other is repeated evolution of a trait in varieties that result from multiple, independent domestication events within the same species.(from reference <ref name="ref1" />)

Revision as of 08:51, 10 June 2014

Rc is a domestication-related gene required for red pericarp in rice (Oryza sativa) and it encodes a basic helix-loop-helix (bHLH) protein that was fine-mapped to an 18.5-kb region on rice chromosome 7 using a cross between Oryza rufipogon (red pericarp) and O. sativa cv Jefferson (white pericarp).

characteristic

Different colors, such as purple, brown, red and white, occur in the pericarp of rice. Here, two genes affecting proanthocyanidin synthesis in red- and brown-colored rice were elucidated, one of the gene is Rc.[1])

Mutation

picture1. Length mutation (from reference [2]).

Clip image0021.gifGrain color has been the target of selection in domesticated Asian rice (as well as other domesticated grains), and the vast majority of O. sativa varieties have white (nonpigmented) grains, compared to the dark red grains that characterize the wild relative and other wild species. The genetic basis of white grains in O. sativa has been pinpointed to loss-of-function mutations in the Rc gene, a regulatory protein in the proanthocyanidin synthesis pathway; these loss-of-function mutations prevent the development of a pigmented pericarp layer. A survey of 337 white pericarp rice cultivars showed that the majority (>97%) have a 14-bp deletion in exon 7 of the gene, resulting in a premature stop codon and a non-functional allele referred to as rc. Our findings for the pericarp color gene, Rc, indicate that remarkably similar genetic mechanisms can be at play. We find that two of the three white pericarp O. glaberrima varieties in our sample harbor a unique point mutation that is predicted to result in a premature stop codon in exon 7 of the Rc gene. Instead, two samples contained a novel point mutation predicted to result in a premature stop-codon in exon 7, hereafter referred to as allele rc-g1 . This mutation is 146 bp upstream of the Rc-s point mutation and 201 bp upstream of the initiation of the 14-bp rc deletion. The third white pericarp O. glaberrima contained no identifiable insertions, deletions, or premature stop-codons, nor does it differ from the red-pericarp O. glaberrima sequences for any predicted amino-acid changes. Comparison of this sample to the most closely related red-pericarp sample showed that they were identical for the sequenced region upstream of the start codon (>1.5 kb) and for the first intron; both of these regions are potentially important as cis-regulatory regions in plants. This study also shows that white pericarps might be achieved by mutations in cis-regulatory regions of Rc or potentially mutations in other genes, a pattern that was not observed in an extensive survey of O. sativa cultivars.(from reference [2]) Crop domestication can serve as a model of plant evolutionary processes. It involves a series of selection events from standing natural variation and newly occurring mutations and combinations of mutations as a result of natural crossings in populations during local adaptation and propagation of plant lines to other cultivation areas.[3]) To determine the subpopulation origin of the Rc mutations,we examined ancestral haplotypes across the Rc coding sequence and promoter region in 103 genetically diverse, redgrained rices. Four rice insertion/deletion polymorphism (RID) and two rice microsatellite (RM) polymorphisms were used to construct haplotypes across the 6.5-kb region containing the Rc gene.[4])

Function

Struture1.jpgRc alleles from the temperate ecotype of weedy lines LD and C9541 could represent the functional alleles most distant to those in group I (Figure 4). All the donor lines of weedy and cultivated rice in both groups were identified as dormant genotypes, or dormancy was associated with the Rc locus in our previous or present research. Thus, phylogenetic and dormancy data suggest that Rc alleles functional for pericarp color most likely also function for seed dormancy.[5])

Expression

RT-PCR experiments confirmed that the Rc gene was expressed in both red- and white-grained rice but that a shortened transcript was present in white varieties.We anticipated that similar expression levels of Rc would be detected in red and white seeds. Our results confirmed this expectation and further demonstrated that the RNA transcript from cv Jefferson contained the 14-bp deletion predicted from the sequence information.(from reference [6])


Nucleotide Polymorphisms

Phylogenetic analysis, supported by comparative mapping in rice and maize (Zea mays), showed that Rc, a positive regulator of proanthocyanidin, is orthologous with INTENSIFIER1, a negative regulator of anthocyanin production in maize, and is not in the same clade as rice bHLH anthocyanin regulators.Mapping of Rc.jpg(from reference [6])

Allele Distribution

Sequencing of the alleles from both mapping parents as well as from two independent genetic stocks of Rc revealed that the dominant red allele differed from the recessive white allele by a 14-bp deletion within exon 6 that knocked out the bHLH domain of the protein.(from reference [6])

Recombination

The rg7.1 locus was originally mapped to a 7.2-Mb region that included the centromere on rice chromosome 7. Given the low frequency of recombination across this region, it was not clear whether positional cloning would be feasible. This study demonstrates that even in regions that are recombinationally repressed, map-based gene isolation offers a viable approach.(from reference [6])

Evolution

Repeated phenotypic evolution can occur at the interspecific level, where it is manifest as the appearance of the same trait in multiple domesticated species, and at the intraspecific level, where the same trait arises multiple times within a single crop. In rice, the opportunity exists to examine the genetic basis of repeated trait evolution at both the interspecific and intraspecific level. At the intraspecific level, repeated phenotypic evolution during domestication can be explored in two contexts; one is the repeated evolution of a trait in varieties resulting from a single domestication event, the other is repeated evolution of a trait in varieties that result from multiple, independent domestication events within the same species.(from reference [2])


Labs working on this gene

  • Department of Biology, Washington University in St. Louis, St. Louis, Missouri, USA
  • Plant Science Department, South Dakota State University, Brookings, South Dakota 57007, †Biosciences Research Laboratory, U.S.
  • Department of Agriculture–Agricultural Research Service, Fargo, North Dakota 58105, ‡Northern Crop Science Laboratory, U.S.
  • Department of Agriculture–Agricultural Research Service, Fargo, North Dakota 58102, §National Institute of Agrobiologica
  • Sciences, Tsukuba, Ibaraki 305-8602, Japan
  • Agricultural College, Yangzhou University, Yangzhou 225008, China
  • Department of Plant Breeding and Genetics, Cornell University, Ithaca, New York 14953-1901
  • Department of Plant Biology, Cornell University, Ithaca, New York 14853
  • Department of Plant Breeding and Genetics, Cornell University, Ithaca, New York, United States of America
  • International Rice Research Institute, Los Ban˜ os, Philippines*
  • Indonesian Center for Agricultural Biotechnology and Genetic Resources Research and Development, Bogor, Indonesia
  • Department of Agronomy, Chungbuk National University, Chongju, Republic of Korea
  • National Institute of Agricultural Biotechnology, Suwon, Republic of Korea
  • Department of Biological Statistics and Computational Biology, Cornell University, Ithaca, New York, United States of America
  • Plant Genome Research Unit, National Institute of Agrobiological Sciences, 2-1-2 Kannondai, Tsukuba, Ibaraki, 305-8602 Japan
  • QTL Genomics Research Center, National Institute of Agrobiological Sciences, 2-1-2 Kannondai, Tsukuba, Ibaraki, 305-8602 Japan
  • Genetic Diversity Department, National Institute of Agrobiological Sciences, Tsukuba, Ibaraki 305-8602, Japan
  • Department of Biological Science and Technology, Tokyo University of Science, Noda, Chiba 278-8510, Japan
  • Research Institute for Biresources, Okayama University, Kurashiki, Okayama 710-0046, Japan
  • National Agricultural Research Center for Kyushu Okinawa Region, National Agriculture and Bio-oriented Research Organization, Nishigoshi, Kikuchi, Kumamoto 861-1192, Japan
  • National Institute for Basic Biology, Okazaki, Aichi 444-8585, Japan
  • Plant Breeding Laboratory, Graduate School of Agriculture, Hokkaido University, Sapporo, Hokkaido 060-8589, Japan

References

  1. Tsutomu Furukawa, Masahiko Maekawa, Tomoyuki Oki, Ikuo Suda, Shigeru Iida, Hiroaki Shimada, Itsuro Takamure, Koh-ichi Kadowaki The Rc and Rd genes are involved in proanthocyanidin synthesis in rice pericarp The Plant Journal, 2006, 49(1): 91-102.
  2. 2.0 2.1 2.2 Xing-You Gu; Michael E. Foley; David P. Horvath; James V. Anderson; Jiuhuan Feng; Lihua Zhang; Chase R. Mowry; Heng Ye; Jeffery C. Suttle; Koh-ichi Kadowaki; Zhongxiang Chen. Association Between Seed Dormancy and Pericarp Color Is Controlled by a Pleiotropic Gene That Regulates Abscisic Acid and Flavonoid Synthesis in Weedy Red Rice;Genetics, 2011, 189(4): 1515-1524.
  3. Saeko Konishi, Kaworu Ebana and Takeshi Izawa Inference of the japonica Rice Domestication Process from the Distribution of Six Functional Nucleotide Polymorphisms of Domestication-Related Genes in Various Landraces and Modern Cultivars Plant and Cell Physiology, 2008, 49(9): 1283-1293.
  4. Megan T. Sweeney; Michael J. Thomson; Yong Gu Cho; Yong Jin Park; Scott H. Williamson; Carlos D. Bustamante; Susan R. McCouch Global Dissemination of a Single Mutation Conferring White Pericarp in Rice PLoS Genetics, 2007, 3(8): 1418-1424.
  5. B. L. GROSS, F. T. STEFFEN, K. M. OLSEN The molecular basis of white pericarps in African domesticated rice: novel mutations at the Rc gene.Journal of Evolutionary Biology, 2010, 23(12): 2747-2753.
  6. 6.0 6.1 6.2 6.3 Megan T. Sweeney, Michael J. Thomson, Bernard E. Pfeil, Susan McCouch Caught Red-Handed: Rc Encodes a Basic Helix-Loop-Helix Protein Conditioning Red Pericarp in Rice The Plant Cell, 2006, 18(2): 283-294.

Structured Information

Gene Name

Os07g0211500

Description

Pericarp Color;red grain color gene

Version

DQ204735.1 GI:78057266

Length

833bp

Definition

Oryza sativa (japonica cultivar-group) cultivar H75 brown pericarp

           and seed coat (Rc) gene, complete cds.
Source

Oryza sativa Japonica Group (Japanese rice)

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

Chromosome 7

Location

Chromosome 7

Sequence Coding Region

{{{CDS}}}

Expression

GEO Profiles:Os07g0211500

Genome Context

{{{GCID}}}

Gene Structure

<gbrowseImage2> name=NC_008397:19856181..19857859 source=RiceChromosome07 preset=GeneLocation </gbrowseImage2>

Coding Sequence

<cdnaseq> 1 atggccggcg gcgaggcgca tgcggcgctg caggcggtgg cgcagagcct ccggtggacc

      61 tacagcctcc tctggcagct ctgcccccac caagggagctcg
     421 ctggtgtggg gggaggggca ctacaacggc gccgtcaaga cgcggaagtc gacggtgatg
     481 cagccgccgc cggcggagga ggaggacgac gccgaccacg cggcgcgcca ccggagccgg
     541 cagctgaggg agctctacga ctggctgcag caggccgggg agaactccag cggcggcgtg
     601 cagacgtcgt cgacgacggc gagccggcgg ccgggggcgg ctctgtcgcc ggaggacctg
     661 acggagacgg agtggttctt cctcatgtcg gcatcctact ccttccctcc cggcatcgggtta
    3421 cctggaaggg catttgcaag gagaggccat gtatggctca ctggagcaaa tgaagttgac
    3481 agcaaagtat tcctaagagc aattcttgcc aagacagttgtg tgcattcctg ttgtcgatgg cgtcctggaa attggaacta cggaaaaggtggaggaag atatgggcct gattcagtat gcaaggggca
    4201 tcttcatgga tcaacatggc atccacatga agcctaccct ctcacagcac tcaacatcca
    4261 acccagtcac ccactgtact catcagcatc caatccaggt tcagatgcaa ctaggtatca
    4321 ccagccaaac aaagtttgat tattcagatg agctcaatgc agatgaggag aatgatgaca
    4381 cagaagaaga gggcatgtca ggttcagaca ctaacaacac tgacactgaa aggaattcag
    4441 gccagctgca acttcaaatg caagaccaac tgaacatggt gagcaatgac caccagacaa
    4501 taccaaataa tgcagtttcc agtgagctaa tgcagtgtga gatgtcagaa gtggtaagag
    4561 atggctgctc aaataatatt ttagaggatg aaatccaaat gctgatggat tgccaaaaca
    4621 gtaattgtca gttaaatttg caagggccag atgagccttg tcactcttgg cattttctct
    4681 gcgaggagtt acaaaatgat taccagccagctactgaaga tcaagtggca tcacctgaaa atacccatta
    4921 cccaaaaaca ctcatgacaa tcctacatta caacacgctg cgacagcaag agatgaacat
    4981 caagaactac ttgccagttt cagagaaatc atcattctcc agatggacta ctcctgaagg
    5041 aagtgatgac aacaagacca tgatcagtcc aggcaccaca cagagaatgc tcaagagcat
    5101 cctgatgatt gttcccagta gtcactgcag ttacagggga gcagaaacac ctgaatcaag
    5161 gggcgggaaa ggcgcaagtg gaacgcgaaa agtcggtgcc atccaaggtg atttcagtgc
    5221 caaccatgtg ctgaaagaga ggagaagaag agagaagctc aatgagaagt tcataattct
    5281 gcgatctttg gtacctttca tgacaaagatggac aaggcgtcga tactaggcga
    6001 cacgatcgag tacgtgaagc agctaaggaa ccgcatacaa gagctcgagt cgtcgtcgtc
    6061 gtcgtcacga gcagccgccc gggcgccatc ggcggcggcc gccgggaggc ggaggaagag
    6121 atccgccgcc gccgccactg ccacggcggc ggaagggatg agcagcagca atggccgcaa
    6181 tggcggcgag gcggcggagg tggtgcaggt gtccatcatc gagagcgacg cgctgctgga
    6241 gctccggtgc ggttgcggcg gcggcggcgg cggtgtggtg ctgctccggg tgatgcaggc
    6301 gatgcaggag ctccagctgg aggtcaccgc cgtccaggcc tcgtgcgccg gtggcgagct
    6361 gctcgccgag ctgcgcgcca aggtcgtcgt tatgatcctg atctgcatga aaatgcaaat
    6421 gcaaatgcaa atgcagaatt aa</cdnaseq>
Protein Sequence

<aaseq>MAGGEAHAALQAVAQSLRWTYSLLWQLCPHQGSSLVWGEGHYNG

                    AVKTRKSTVMQPPPAEEEDDADHAARHRSRQLRELYDWLQQAGENSSGGVQTSSTTAS
                    RRPGAALSPEDLTETEWFFLMSASYSFPPGIGLPGRAFARRGHVWLTGANEVDSKVFL
                    RAILAKTVVCIPVVDGVLEIGTTEKVEEDMGLIQYARGIFMDQHGIHMKPTLSQHSTS
                    NPVTHCTHQHPIQVQMQLGITSQTKFDYSDELNADEENDDTEEEGMSGSDTNNTDTER
                    NSGQLQLQMQDQLNMVSNDHQTIPNNAVSSELMQCEMSEVVRDGCSNNILEDEIQMLM
                    DCQNSNCQLNLQGPDEPCHSWHFLCEELQNDYQPATEDQVASPENTHYPKTLMTILHY
                    NTLRQQEMNIKNYLPVSEKSSFSRWTTPEGSDDNKTMISPGTTQRMLKSILMIVPSSH
                    CSYRGAETPESRGGKGASGTRKVGAIQGDFSANHVLKERRRREKLNEKFIILRSLVPF
                    MTKMDKASILGDTIEYVKQLRNRIQELESSSSSSRAAARAPSAAAAGRRRKRSAAAAT
                    ATAAEGMSSSNGRNGGEAAEVVQVSIIESDALLELRCGCGGGGGGVVLLRVMQAMQEL
                    QLEVTAVQASCAGGELLAELRAKVVVMILICMKMQMQMQMQN"</aaseq>
Gene Sequence

<dnaseqindica>1321..1386#668..1152#450..531# 1 atggccggcg gcgaggcgca tgcggcgctg caggcggtgg cgcagagcct ccggtggacc

      61 tacagcctcc tctggcagct ctgcccccac caagggtacc taccctacct acctacgaca
     121 cgatgcacag tgttcatcca tggccggcca tggcggatcg tcgtcgttgt cgatgatcat
     181 cgaaggaagc tagaggatat ggctcaatac tttgataata tatatactga tctctccgta
     241 caacaaaaat ataaaaattc tagctagtat cgaatgagac atatgctatg ctagtactac
     301 gaatctaaaa agatgtacat attttgattc gtattattag gatatatcac gagtttttat
     361 attttgagac ggatgtaata attctgaatt tagttgtgat ccatggcatg caggagctcg
     421 ctggtgtggg gggaggggca ctacaacggc gccgtcaaga cgcggaagtc gacggtgatg
     481 cagccgccgc cggcggagga ggaggacgac gccgaccacg cggcgcgcca ccggagccgg
     541 cagctgaggg agctctacga ctggctgcag caggccgggg agaactccag cggcggcgtg
     601 cagacgtcgt cgacgacggc gagccggcgg ccgggggcgg ctctgtcgcc ggaggacctg
     661 acggagacgg agtggttctt cctcatgtcg gcatcctact ccttccctcc cggcatcggg
     721 tatataataa aaaatataga tataaatatt taagcatgca tgcataaatt aaaccacact
     781 tcttgttacg tgttcttggc aaaatgatga acaattacca ctaattaatt ggagccagaa
     841 accctaaaga tttacccacc tggttaatta atcggtgtgt tgatccacgc atgcatgcat
     901 gcagaaaatc aagatcagga tagctccttt tcttttgcag gttaattagc tagatcttca
     961 cgtataatta gctagctaga ttttaaaata taatttattc aatttgattt atgattttta
    1021 ttttttattt caaatagata caactgtata caaaatttta ttttggtaca tacctccgat
    1081 ccaactacat cagaggtaaa aaaaaaatta aaccgttgga attgattaga acaagatcgt
    1141 gcggtcaaat tatatcataa ctaacttttc tgattctcta aagcatagag atgtatatat
    1201 acatcgtatt attaggctct atatttcctg attaacacta gatgcatata taattttgat
    1261 agtcaaaata tacttttgat aggctctaaa gaaaaactta ataacatgta ctccctccat
    1321 atacttttga tagtcatatt tcatcttgac acacagatca agtataagta attctactta
    1381 tcatccattt aaacacgcta ctagttattc ctcataaaca agcgattcat taatatttac
    1441 atttctcgat gcttgtgtag ccaatattgt gtggaagaat ggaatgtcat taagaggata
    1501 ggttgttgga ttgaaatatg cctatcaaaa ataaattttt agatttgaaa atatgcctat
    1561 caaaagtaga tggagggagt attaattaat gtgaatttcc aatcctactg ttgtgatatt
    1621 aggctttgta ccttcttgtc caggaggtat atatatggct cttttaagga tgggagaaaa
    1681 tatcatcttt aatacaacta tatatggctt ttgtttgata aatacaactt ttattttgta
    1741 tgaatacaaa tatattgata aatatccacc attataatcc taacccatta ggatcatatg
    1801 gtgtatattt ttttaactat ttgtttttta taaattaata ttaagagatc acaataaaaa
    1861 tatagtatta tgaaagtact cttaacaaca tatccaatga taaaattatt attattacaa
    1921 aatatagtgg tcaaattgta tagaattcaa tagcctgatt ttatgacgtc aagtaaatta
    1981 aataaagaat gaaggtagtg ctagagtgat caaacaatat ctctcctaaa atatgtccta
    2041 taagttttac tccataaatc caagggtcaa aagttgttgg gttatttttt tagataataa
    2101 catactaccc cttttcaaaa tgtatgattc tattgacttt ttgcacaaca tttaaccatt
    2161 tgtcatatta aaaattagta taaacatcta aaaatataag ttacaattat attttatttg
    2221 atgataaaac aactcacaac aaaataaata atatttatat aatctttttg gaataaaacg
    2281 aatgatcaaa cattattcaa aaagtcaatg gtatagtacg ttttgaaatt gatagactat
    2341 gagagcaaaa ttttgagata acatggaaaa ttatcctctt agacattgca ctgtgtaata
    2401 attaataata atgaatgaaa ggctaagact tttcttccac cttatataag tggttgaata
    2461 tatagcaatc acatcattac atgattttgt aaccaaccgt ctctatagct ccgatacagt
    2521 gctagtttca catcgtaata attaaagagt ataataataa atcgaggtgt acttctcatc
    2581 gatgaagtga tgtgccgctt agctaaatta aactcgtatg cgaaaaatca gtatatgtcc
    2641 ggttaatttc taagagagag attgagagag aataattgcg cccctccaaa tccccctctt
    2701 ggacgttagg gagctatata gacggtattg ctaagtgcga tgtgtacata acgtacctgt
    2761 cgtaggaaca tttctcatcc aaattaagta gtaatgcatg gcatgaaatc catttttgta
    2821 ttttgcatgg caaagaatga caacaaggaa tacactagct agccctgccc tttttcaatt
    2881 taatttaaca tcaaacttag ttattgtatt tcttttgtca gaatagcatg cattgcatac
    2941 tctttaaaaa taattaatta gtgtatttta ctagtcttac aaaagtatca agagagacaa
    3001 ctaattatag ttgggagaca ccaaacttgt ttttaataat gacaattaaa accctacctc
    3061 tacatccaac atagacgtac atagtccgaa ggcgccaaat atttgtacat ttagctacca
    3121 gatttcagta cgagttctca cattataatt ttgatttttt tatttttttt ataaacaatc
    3181 tggtaccctt ttatgtctgg aaggaaaaaa aaaatctaaa ttgcaacatt ttagtcggtg
    3241 agaatggtac tctgtcctag ctactttcta cacatgagag agagagagag agagagagag
    3301 agagccttta attgcccttg cccatgcatc tttctttgca cacatgtatg cttttcacat
    3361 tgtcatgagg agagaacttg ttaagttgca cacatgtgtg ctttgcatgt cttcaggtta
    3421 cctggaaggg catttgcaag gagaggccat gtatggctca ctggagcaaa tgaagttgac
    3481 agcaaagtat tcctaagagc aattcttgcc aaggttcagc catcaccttc tcttacctat
    3541 ttttcactct gaatgccaac agtgctttgc acattgtagt ctgtttgcag actgcaaatg
    3601 atgaccataa tcagatcaga aaataaaata atattatata ctttttgagc cagctagcaa
    3661 gaatatgtaa caataattct cctttttttt tcttgttctt ttccctgatg tggtgcataa
    3721 caaataacca aactgatgaa tggcagagtg ctggtatcca ggtatttgcc tctaaaagta
    3781 gctacacgtt tactatgaaa ttttgtggct tttgttcatc tttggatgca gtggccatta
    3841 tctaaaaact atgaatttcc agactgcagt ttttatctaa ttttgtgact ttgtacatca
    3901 gacagttgtg tgcattcctg ttgtcgatgg cgtcctggaa attggaacta cggaaaaggt
    3961 gatttcgtat attatcagct gacaatctaa ttatatgggc catataatta agtataaatc
    4021 aaaatacctc ataatatatt ataaagtatc taatgtgatt atgtgaatat tggctatttc
    4081 aatgtaattt gatatatgaa actgataatc ctctgaaact ccgtaaggat caaactaatc
    4141 aaaatgtata tattttcaag gtggaggaag atatgggcct gattcagtat gcaaggggca
    4201 tcttcatgga tcaacatggc atccacatga agcctaccct ctcacagcac tcaacatcca
    4261 acccagtcac ccactgtact catcagcatc caatccaggt tcagatgcaa ctaggtatca
    4321 ccagccaaac aaagtttgat tattcagatg agctcaatgc agatgaggag aatgatgaca
    4381 cagaagaaga gggcatgtca ggttcagaca ctaacaacac tgacactgaa aggaattcag
    4441 gccagctgca acttcaaatg caagaccaac tgaacatggt gagcaatgac caccagacaa
    4501 taccaaataa tgcagtttcc agtgagctaa tgcagtgtga gatgtcagaa gtggtaagag
    4561 atggctgctc aaataatatt ttagaggatg aaatccaaat gctgatggat tgccaaaaca
    4621 gtaattgtca gttaaatttg caagggccag atgagccttg tcactcttgg cattttctct
    4681 gcgaggagtt acaaaatgat taccagccag gtattacatt tgagaagata atccttcaaa
    4741 agcacccttg ttccaaaaat atatatttgt actcttcaca caagcactgc catttttttt
    4801 cttttttgca tacatcctca attcttgcat ttcttttcca tatatttgat acaactgtct
    4861 ccatttccct tctgtcacag ctactgaaga tcaagtggca tcacctgaaa atacccatta
    4921 cccaaaaaca ctcatgacaa tcctacatta caacacgctg cgacagcaag agatgaacat
    4981 caagaactac ttgccagttt cagagaaatc atcattctcc agatggacta ctcctgaagg
    5041 aagtgatgac aacaagacca tgatcagtcc aggcaccaca cagagaatgc tcaagagcat
    5101 cctgatgatt gttcccagta gtcactgcag ttacagggga gcagaaacac ctgaatcaag
    5161 gggcgggaaa ggcgcaagtg gaacgcgaaa agtcggtgcc atccaaggtg atttcagtgc
    5221 caaccatgtg ctgaaagaga ggagaagaag agagaagctc aatgagaagt tcataattct
    5281 gcgatctttg gtacctttca tgacaaaggt aattaagtac tccctctatt tctataaagc
    5341 cgtatttgac tagttatctt atttagaaag tatgtgcaaa tatgtaaaat ataagtcata
    5401 cttaaagaac ttttaatgtt attaaataat aagtcacacc aaaaataaaa catatatatt
    5461 tttaataaga caaatgatta aatgtatata taaaaattaa tagcgtcaca tattttaaaa
    5521 tagaggggta tttaagtacc cacaggatca tcaaaattca gttatctttt cttaagcctc
    5581 taacgaacat tggaagatcc tcactaatgg cagcatgaat ctagggttca ctatttcgga
    5641 atgcaaaata tgttttaccg ggcatccgat ttttaaaaaa ttcagaatga agaaaattga
    5701 atctttttta tggatttgaa taaatcttga taaattcgaa aaaatttccg aacttttggc
    5761 cagaagtgaa tcctacccgt atccaccggt aataaaccta aatttttggg agtaatgaat
    5821 taatgttata tataatccat gaattatata gttccaaact actccgtaac aaattttcag
    5881 gagtagtgaa attaatatta ttacaatctc agaaaaaaat ggcagaaaca attaatctgt
    5941 tttcaattat taattaattt gtttttgtgt ccagatggac aaggcgtcga tactaggcga
    6001 cacgatcgag tacgtgaagc agctaaggaa ccgcatacaa gagctcgagt cgtcgtcgtc
    6061 gtcgtcacga gcagccgccc gggcgccatc ggcggcggcc gccgggaggc ggaggaagag
    6121 atccgccgcc gccgccactg ccacggcggc ggaagggatg agcagcagca atggccgcaa
    6181 tggcggcgag gcggcggagg tggtgcaggt gtccatcatc gagagcgacg cgctgctgga
    6241 gctccggtgc ggttgcggcg gcggcggcgg cggtgtggtg ctgctccggg tgatgcaggc
    6301 gatgcaggag ctccagctgg aggtcaccgc cgtccaggcc tcgtgcgccg gtggcgagct
    6361 gctcgccgag ctgcgcgcca aggtcgtcgt tatgatcctg atctgcatga aaatgcaaat
    6421 gcaaatgcaa atgcagaatt aa

</dnaseqindica>

External Link(s)

NCBI Gene:Os07g0211500, [1]