Difference between revisions of "Os11g0225300"
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recognized by the product of the so far uncharacterized Pi-CO39R gene (Ribot et al., 2013).<ref name="ref3" /> | recognized by the product of the so far uncharacterized Pi-CO39R gene (Ribot et al., 2013).<ref name="ref3" /> | ||
The molecular mechanism of M. oryzae Avr protein recognition has only been investigated in the case of AVR-Pita and AVR-Pik (Jia et al., 2000; Kanzaki et al., 2012). AVR-Pita is recognized through direct binding to the Pi-ta C-terminal LRD domain, whereas AVR-Pik specifically associates with an N-terminal domain of Pik-1, including the CC domain and additional unclassified sequences upstream of the NB domain.Hence, those examples illustrate two cases of direct recognitionthat seem to implicate different R protein domains and different mechanisms.<ref name="ref4" /> | The molecular mechanism of M. oryzae Avr protein recognition has only been investigated in the case of AVR-Pita and AVR-Pik (Jia et al., 2000; Kanzaki et al., 2012). AVR-Pita is recognized through direct binding to the Pi-ta C-terminal LRD domain, whereas AVR-Pik specifically associates with an N-terminal domain of Pik-1, including the CC domain and additional unclassified sequences upstream of the NB domain.Hence, those examples illustrate two cases of direct recognitionthat seem to implicate different R protein domains and different mechanisms.<ref name="ref4" /> | ||
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| + | [[File:AVR1-CO39 Interacts Physically with RGA5.jpg|right|thumb|150px|''Semidwarf VS. normal-type rice plants at ripening (from reference <ref name="ref3" />).'']] | ||
===Evolution=== | ===Evolution=== | ||
Revision as of 04:38, 3 June 2014
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Contents
Annotated Information
Function
Plant resistance to microbial pathogens is a complex process relying on two major levels of resistance controlled by distinct types of plant receptors (Jones and Dangl, 2006; Dodds and Rathjen, 2010). The first line of plant defense is activated by plasma membrane proteins called pattern recognition receptors,which perceive conserved microbial molecules called pathogenassociated molecular patterns (PAMPs). Adapted plant pathogens are able to bypass this PAMP-triggered immunity by producing secreted effectors that act inside or outside the host cell and manipulate key components of plant defense (Jones and Dangl, 2006). The second layer of plant immunity relies on the specific recognition of certain pathogen-derived effectors called Avirulence (Avr) proteins by so-called plant resistance (R) proteins. This effector-triggered immunity (ETI) gives rise to stronger and faster defense responses than PAMP-triggered immunity and often involves a form of localized programmed cell death called the hypersensitive response (HR) (Dodds and Rathjen, 2010). The largest class of R proteins belongs to the conserved family of NB-LRR proteins (Tameling and Takken,2007). They contain a central nucleotide binding (NB) domain,also known as the NB-ARC (for NB adaptor shared by Apaf-1,certain R proteins, and CED-4) domain, and a C-terminal leucinerich repeat (LRR) domain. In monocot R proteins, the LRR repeatmotif is often not conserved (Bai et al., 2002) and in those cases,the domain is called leucine-rich domain (Monosi et al., 2004;Zhou et al., 2004). NB-LRR proteins are further subdivided according to their N-terminal domain into two major subclasses(Meyers et al., 1999; Pan et al., 2000). Proteins of the TIR-NB-LRR class possess an N-terminal Toll Interleukin-1 (TIR) domain,whereas CC-NB-LRR class proteins harbor a structured coiledcoil(CC) domain. Both N-terminal domains seem to be involved in R protein homodimerization and in the activation of defensesignaling (Bernoux et al., 2011; Maekawa et al., 2011). In the absence of the Avr protein, R proteins are maintained in an inactive conformation to avoid inappropriate defense activationand cell death (Takken and Goverse, 2012).[1] [2]
Expression
Seven Avr genes from M. oryzae have been cloned. Except ACE1 and AVR-Pita, which encode an enzyme involved in the synthesis of a secondary metabolite (Böhnert et al., 2004) and a putative metalloprotease (Orbach et al., 2000), respectively,Avr genes from the rice blast fungus encode small secreted proteins of unknown function. Experimental evidence indicates that recognition of AVR-Pita, AVR-Pia, and AVR-Pik/km/kp occurs inside host cells by their corresponding cytoplasmic R
proteins (Jia et al., 2000; Yoshida et al., 2009; Kanzaki et al.,2012). Recently, we characterized molecularly the AVR1-CO39gene and demonstrated that it encodes a small secreted protein,expressed specifically during infection (Ribot et al., 2013). AVR1-CO39 is translocated inside the cytoplasm of rice cells where it is recognized by the product of the so far uncharacterized Pi-CO39R gene (Ribot et al., 2013).[3] The molecular mechanism of M. oryzae Avr protein recognition has only been investigated in the case of AVR-Pita and AVR-Pik (Jia et al., 2000; Kanzaki et al., 2012). AVR-Pita is recognized through direct binding to the Pi-ta C-terminal LRD domain, whereas AVR-Pik specifically associates with an N-terminal domain of Pik-1, including the CC domain and additional unclassified sequences upstream of the NB domain.Hence, those examples illustrate two cases of direct recognitionthat seem to implicate different R protein domains and different mechanisms.[4]
Evolution
Please input evolution information here.
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Labs working on this gene
- INRA, UMR 385 Biologie et Génétique des Interactions Plante-Parasite, F-34398 Montpellier, France
- CIRAD, UMR Biologie et Génétique des Interactions Plante-Parasite, F-34398 Montpellier, France
- CNRS, Plateforme Imagerie-Microscopie, Fédération de Recherche FR3450, 31326 Castanet-Tolosan, France
- INRA, UMR 441 Laboratoire des Interactions Plantes-Microorganismes, F-31326 Castanet-Tolosan, France
- CNRS, UMR 2594 Laboratoire des Interactions Plantes-Microorganismes, F-31326 Castanet-Tolosan, France
- Iwate Biotechnology Research Center, Kitakami, Iwate 024-0003, Japan
References
- ↑ Stella Cesari;Gaëtan Thilliez;Cécile Ribot, et al. (2013) The Rice Resistance Protein Pair RGA4/RGA5 Recognizes the Magnaporthe oryzae Effectors AVR-Pia and AVR1-CO39 by Direct Binding.The Plant Cell, 25(4): 1463-1481.
- ↑ Altschul, S.F., Madden, T.L., Schäffer, A.A., Zhang, J., Zhang, Z.,Miller, W., and Lipman, D.J. (1997). Gapped BLAST and PSIBLAST:A new generation of protein database search programs.Nucleic Acids Res. 25: 3389–3402.
- ↑ 3.0 3.1 Ashikawa, I., Hayashi, N., Yamane, H., Kanamori, H., Wu, J.,Matsumoto, T., Ono, K., and Yano, M. (2008). Two adjacent nucleotide-binding site-leucine-rich repeat class genes are required toconfer Pikm-specific rice blast resistance. Genetics 180: 2267–2276.
- ↑ Bai, J., Pennill, L.A., Ning, J., Lee, S.W., Ramalingam, J., Webb,C.A., Zhao, B., Sun, Q., Nelson, J.C., Leach, J.E., and Hulbert,S.H. (2002). Diversity in nucleotide binding site-leucine-rich repeatgenes in cereals. Genome Res. 12: 1871–1884.
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Structured Information
| Gene Name |
Os11g0225300 |
|---|---|
| Description |
Disease resistance protein family protein |
| Version |
NM_001074069.1 GI:115484786 GeneID:4350112 |
| Length |
7166 bp |
| Definition |
Oryza sativa Japonica Group Os11g0225300, 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 | |
| Location |
Chromosome 11:6537450..6544615 |
| Sequence Coding Region |
6537456..6538285,6539404..6541536,6543349..6544291 |
| Expression | |
| Genome Context |
<gbrowseImage1> name=NC_008404:6537450..6544615 source=RiceChromosome11 preset=GeneLocation </gbrowseImage1> |
| Gene Structure |
<gbrowseImage2> name=NC_008404:6537450..6544615 source=RiceChromosome11 preset=GeneLocation </gbrowseImage2> |
| Coding Sequence |
<cdnaseq>atggatgctccggtgagtttctccctaggtgccatgggccctctcctcaggaagctcgattcgcttccggtggctcccgaaatccggctgcctgagccactgaaggacggaatcgagctcctcaaggaagatctagaagaaatcggtgccgccctcgtggagcagtccatggtagattccccaagccacagggcaagatactggatggatgaggtgcgtgacctctcctaccacatcgaggactgcatcgacaccatgttttcgatgcgctgcggcggcgacgacggcaagcccagatccgttcgccgccacaaggttggccgtgtgaaggttgacggtttctccaagacgcagaagccgtgcacgaggctcgccaggatcgcggagctcagggctctggtgcgggaggccagcgagcgccacgaaaggtaccagcttggtgacggccgcgcctcgagctcgagctcgagctctcaccgcgtgttcactgcgcacggccaggttcctgcgccgtgcagaaatctggttggcatggacgaaccgaagacgaagctcaccaatatgctcaccgacgaagcagagctgcatatgaaagtggtatgcattcttggatcagccggtattggtaagaccacgcttgctgaacaagtataccgtaaacttagatggcaattcgattgccatgcttttgtgcgggcatcacgaaaacctgacatgaggaggcttcttggggccatcctctctcaagttcagctgcgtatacggatatctgataccagcacggtgcagagccttattgataatctctgggagtatctacagaaaaagaggtattttattgtaattgatgaattatatgaaacagcaacatgggatatcattaccagtgcttttccagaggataataattgcagtaggataatgactacggctggaattgaaggtgtagctctcgaatgctgcagttatcattctgtcaatattttcaagatgatacctcttggccttgatgattctgcaaaattattctttaatagagtttttggatcagagcaacaatgtccttatgaattgaatgaagtttcatataggattacagcaaaatgtggtggattgccactagcagtcattattatagccggtcttttagcaagcctgccgtgcaaaacagagctgtggtacaatatagatggatgtttatgttccagtgtgactacagatattgatttggatgagatactgaaggaaatcataagtcttggctacgacaatcttccacattatttgaagacatgtctgttgtatcttagtttatattcggagggtttcataatttggacggctgacttactgaagcaatggatatctgaaggtttcatcgctgtaatagatggagaagacatagaggaagtcgctgagagctatttctataaccttgttaacaggggaatgatccaaagtgtgaaaactaagtacaataatcaggtgttgtgtactgtgcaccacactgtatttgatcttattattcacaagtccaaagaagagaaatttatctccgcaatagattactctcaaactatgccaggaaattctttagaggctcgtcgattgtcattccactttagcaataccagatatgcaactgaagttgcaggtattacactgtcacaagttcgttcgtttgcctttcttggacttctgaagtgtatgccttctattatggaatttaagcttcttcgggttctaattcttgaattttggggagataaccatggatgcatgagttttaacgttgcaagaatttgtagattgtttcagcttagatatttgaagatctcaagccaaataataatagaattaccagcccagatccgggggctaaaatacttggaaacacttgaaatagatgcaagagtaactgcgattccatctgatattattcatctccgaagtttgttgcatctatattttcaagatgggatcgttctacctgatggcattggctgcatcagatctttgcgtacacttaagtattttgatcttggaagtaactctgaagagaatatacggagtcttggtcagctgacgaacttacgggatcttcatctcacttgttctgcaccaaaatctaatcagcaggcaaagagaaacctcgtaattctagcctcttatactgggaaactcggtaacctcaaatctgtcaaattttctcctggtgattcaggcatggatatttcctttttgttttatggaattggtatttcggttgatcgttccagaacagcgtcctctcttcctttctctgttaggacattagagttgccatccatttgcatatttgctagacttcctgactggattggtcaactccgtaaactccacacattgaatcttgctgttagagaactgatagaaaatgatattgatagtcttgcaggactgcctgacctcattgttctctcaatgcatatcatgaaagctcctatggaaaggattgtcttcaacagaaaagcatttccagttctcaagtatttcaagtttatatgtggcacactgcgcatggcttttcaggcaggagccatggccaatcttcacaggctcaagctaggtttcaatgcccataaaggggagaagtatgacaatattcttgtaggcattgagcacctattaaacctcaagaagattgctgttcgaattggaggagctgctgaagccaaggagtctgacaggatggctgcagaggctgcgttgaaagaagccattagaaagcatctaatgtttcttgatgatctcgacatagcaagggtagaatgtgttaaggaagagtataagtgtattaagaaaaaacacaagatcaagattgaagattcaattagcgaaaaaaatggggactcaaaaaagcagcattctgtcgaaaagaaagcagtgtggggcaaaactatgaagaatattgctgatagtggcgtgtttcctgaagattatactatgtcaagggagcaacgggtcgcagaaggcttcgtagtgggcattgaaaagtgcagggcagaggatgcggccgagcgaataatcaggaatgtgcctgtggactatgatgggcttggccaggtgagcacatccaagatacaggaccacttgcctgaacttgctcctcgtgctgtgcaaaacgagaagttcggctcttcaaatgatcttagcataatgatacaaatcaacaagtatgctcgtctgccatcttatgagtggagggacactgatatatccaaactgaatttccggcttctccgcgccccgatgttacttgaagctgtaacggcccgttgtcacttgttggacttaatcctcattggttcaaacaacattactgttcttgaccttggacgtcccacaatcaccaaactgccggcatctatagaatgtctacctaatttacggtacttgaggttgcagggcacgcaactgaagtcgctgtcggaggttattgtgaagatgcccaccatcagggggttggacatcaagaataccaagacagaggagctaccacaaggcattttgaggatgaagaagttgagtcacctgagtatgggggagaagcagaaaaatattcaagtattcatggaaaaaatgcagactctggctgagaccgtccaagacagcgatgacttgtccgacgaaacagagggaatagccgacgacgaaggcgagttctctacaagggccaacgcgtcaacaccgaaggtggatgaggacgaggtggaccggagggccaataacttcatcgccaggttcaggaagcagatcacgatcaggaattcaggattcgccaaaaaggagagcagcatcgacgaacgactgtggatacgagatctcgatgagtgccaactctccaagagaggtggacgattttaa</cdnaseq> |
| Protein Sequence |
<aaseq>MDAPVSFSLGAMGPLLRKLDSLPVAPEIRLPEPLKDGIELLKED LEEIGAALVEQSMVDSPSHRARYWMDEVRDLSYHIEDCIDTMFSMRCGGDDGKPRSVR RHKVGRVKVDGFSKTQKPCTRLARIAELRALVREASERHERYQLGDGRASSSSSSSHR VFTAHGQVPAPCRNLVGMDEPKTKLTNMLTDEAELHMKVVCILGSAGIGKTTLAEQVY RKLRWQFDCHAFVRASRKPDMRRLLGAILSQVQLRIRISDTSTVQSLIDNLWEYLQKK RYFIVIDELYETATWDIITSAFPEDNNCSRIMTTAGIEGVALECCSYHSVNIFKMIPL GLDDSAKLFFNRVFGSEQQCPYELNEVSYRITAKCGGLPLAVIIIAGLLASLPCKTEL WYNIDGCLCSSVTTDIDLDEILKEIISLGYDNLPHYLKTCLLYLSLYSEGFIIWTADL LKQWISEGFIAVIDGEDIEEVAESYFYNLVNRGMIQSVKTKYNNQVLCTVHHTVFDLI IHKSKEEKFISAIDYSQTMPGNSLEARRLSFHFSNTRYATEVAGITLSQVRSFAFLGL LKCMPSIMEFKLLRVLILEFWGDNHGCMSFNVARICRLFQLRYLKISSQIIIELPAQI RGLKYLETLEIDARVTAIPSDIIHLRSLLHLYFQDGIVLPDGIGCIRSLRTLKYFDLG SNSEENIRSLGQLTNLRDLHLTCSAPKSNQQAKRNLVILASYTGKLGNLKSVKFSPGD SGMDISFLFYGIGISVDRSRTASSLPFSVRTLELPSICIFARLPDWIGQLRKLHTLNL AVRELIENDIDSLAGLPDLIVLSMHIMKAPMERIVFNRKAFPVLKYFKFICGTLRMAF QAGAMANLHRLKLGFNAHKGEKYDNILVGIEHLLNLKKIAVRIGGAAEAKESDRMAAE AALKEAIRKHLMFLDDLDIARVECVKEEYKCIKKKHKIKIEDSISEKNGDSKKQHSVE KKAVWGKTMKNIADSGVFPEDYTMSREQRVAEGFVVGIEKCRAEDAAERIIRNVPVDY DGLGQVSTSKIQDHLPELAPRAVQNEKFGSSNDLSIMIQINKYARLPSYEWRDTDISK LNFRLLRAPMLLEAVTARCHLLDLILIGSNNITVLDLGRPTITKLPASIECLPNLRYL RLQGTQLKSLSEVIVKMPTIRGLDIKNTKTEELPQGILRMKKLSHLSMGEKQKNIQVF MEKMQTLAETVQDSDDLSDETEGIADDEGEFSTRANASTPKVDEDEVDRRANNFIARF RKQITIRNSGFAKKESSIDERLWIRDLDECQLSKRGGRF</aaseq> |
| Gene Sequence |
<dnaseqindica>7..836#1955..4087#5900..6842#ggcggcatggatgctccggtgagtttctccctaggtgccatgggccctctcctcaggaagctcgattcgcttccggtggctcccgaaatccggctgcctgagccactgaaggacggaatcgagctcctcaaggaagatctagaagaaatcggtgccgccctcgtggagcagtccatggtagattccccaagccacagggcaagatactggatggatgaggtgcgtgacctctcctaccacatcgaggactgcatcgacaccatgttttcgatgcgctgcggcggcgacgacggcaagcccagatccgttcgccgccacaaggttggccgtgtgaaggttgacggtttctccaagacgcagaagccgtgcacgaggctcgccaggatcgcggagctcagggctctggtgcgggaggccagcgagcgccacgaaaggtaccagcttggtgacggccgcgcctcgagctcgagctcgagctctcaccgcgtgttcactgcgcacggccaggttcctgcgccgtgcagaaatctggttggcatggacgaaccgaagacgaagctcaccaatatgctcaccgacgaagcagagctgcatatgaaagtggtatgcattcttggatcagccggtattggtaagaccacgcttgctgaacaagtataccgtaaacttagatggcaattcgattgccatgcttttgtgcgggcatcacgaaaacctgacatgaggaggcttcttggggccatcctctctcaagttcagctgcgtatacggatatctgataccagcacggtgcagagccttattgataatctctgggagtatctacagaaaaagaggtaattaattaacgctatattcactgctttttaatactagttcagctactgagcatcaccaatagactctcaaaattctattcccaaaaatctgttattgggcgttctaaactgattttggggtgttgaatgtattcctccttccaacaaattggcttaattaactgcctaaaattaaggacttggcccatgtaaaaactattaacagaaatcatgctccccatcatccatcatccagcgtgatcacatctgcatgatcggccacgcctccctccgcgatcactcccgccatgattttgggagtcaaccctctcccctagaagagtagagcaagacatacgggaatgggagctgcaaaattttggtagacctttcggatattgttggagcaagatttttacaaaaaaaattgtaaattttggtattgggaatgattttgggagactcttgtgaggctatctccaacccttcatgtcattagcatgaggggactcagtgtccatagcattaagtatattgtcacatcggaaaaaatctgaggtggcaagagagttaatgaggagagagatgaaaagatgaagaaaccatctcttcacaacaaccaagaatgaaaacaagtgataggttcataaaaggagagagaagaaagatgattggatatatagtgtctatatgacatagcaagtatggaaactactttgatagtttctaggttggggatggcctgatgctgcacattgaacaattgctgttccctccgtcacattgaataaggcgcgcatgcattttaaaattgaactctctaaatatctgatcaataatattttgttataaattaacttttatttgacaaaaataatatcattggattgacatttaaatttactttgatataattgtgattttgttgttacaaatatcatagtattttagatattcaaaatcaaagatttgttttaaaaaccataccaagtttgactgcgccggataaagggggtaatttagaaaataaataaataaataaatatattatcctgataaacttcctttcctttttgagagagacctgatacactttgttcttttcccagagaatataatttaattatactttgcaggtattttattgtaattgatgaattatatgaaacagcaacatgggatatcattaccagtgcttttccagaggataataattgcagtaggataatgactacggctggaattgaaggtgtagctctcgaatgctgcagttatcattctgtcaatattttcaagatgatacctcttggccttgatgattctgcaaaattattctttaatagagtttttggatcagagcaacaatgtccttatgaattgaatgaagtttcatataggattacagcaaaatgtggtggattgccactagcagtcattattatagccggtcttttagcaagcctgccgtgcaaaacagagctgtggtacaatatagatggatgtttatgttccagtgtgactacagatattgatttggatgagatactgaaggaaatcataagtcttggctacgacaatcttccacattatttgaagacatgtctgttgtatcttagtttatattcggagggtttcataatttggacggctgacttactgaagcaatggatatctgaaggtttcatcgctgtaatagatggagaagacatagaggaagtcgctgagagctatttctataaccttgttaacaggggaatgatccaaagtgtgaaaactaagtacaataatcaggtgttgtgtactgtgcaccacactgtatttgatcttattattcacaagtccaaagaagagaaatttatctccgcaatagattactctcaaactatgccaggaaattctttagaggctcgtcgattgtcattccactttagcaataccagatatgcaactgaagttgcaggtattacactgtcacaagttcgttcgtttgcctttcttggacttctgaagtgtatgccttctattatggaatttaagcttcttcgggttctaattcttgaattttggggagataaccatggatgcatgagttttaacgttgcaagaatttgtagattgtttcagcttagatatttgaagatctcaagccaaataataatagaattaccagcccagatccgggggctaaaatacttggaaacacttgaaatagatgcaagagtaactgcgattccatctgatattattcatctccgaagtttgttgcatctatattttcaagatgggatcgttctacctgatggcattggctgcatcagatctttgcgtacacttaagtattttgatcttggaagtaactctgaagagaatatacggagtcttggtcagctgacgaacttacgggatcttcatctcacttgttctgcaccaaaatctaatcagcaggcaaagagaaacctcgtaattctagcctcttatactgggaaactcggtaacctcaaatctgtcaaattttctcctggtgattcaggcatggatatttcctttttgttttatggaattggtatttcggttgatcgttccagaacagcgtcctctcttcctttctctgttaggacattagagttgccatccatttgcatatttgctagacttcctgactggattggtcaactccgtaaactccacacattgaatcttgctgttagagaactgatagaaaatgatattgatagtcttgcaggactgcctgacctcattgttctctcaatgcatatcatgaaagctcctatggaaaggattgtcttcaacagaaaagcatttccagttctcaagtatttcaagtttatatgtggcacactgcgcatggcttttcaggcaggagccatggccaatcttcacaggctcaagctaggtttcaatgcccataaaggggagaagtatgacaatattcttgtaggcattgagcacctattaaacctcaagaagattgctgttcgaattggaggagctgctgaagccaaggagtctgacaggatggctgcagaggctgcgttgaaagaagccattagaaagcatctaatgtttcttgatgatctcgacatagcaagggtagaatgtgttaaggaagagtataagtgtattaagaaaaaacacaagatcaagattgaagattcaattagcgaaaaaaatggggactcaaaaaagcagcattctgtcgaaaagaaagcagtgtggggcaaaactatgaagaatattgctgatagtgggtatgttctttcctattatttcagcctctacattttccaatatatattgtttagaaataaatgattactacaacaataaccttatagtaaaacgctaaaacacagttgtatcaaaaggaatctttacaaaggtacatctatgctctttgaaaaggataagatgggatgctctatactaattggagcctctactagaagctttcacattaaccttagaaggcaaaagactggaaattcccacattaacaaaaaagaaaactaaaagcattaatctttgaaagatcaaatggtatgcgccaatgtatcaaatctaaggttgatatcttccaccacagaatatgtgcatttatcaaatttggaatcaataaatacctagttatacttggtttggacactcgagtatagtattttgatgaataataattgaaacattcagcattaacaaatattgagttttactcggtctagagtctcactagcttattttttatgattttattttataattccgaaattcagaattgataaatactgtgttgtactaggtttgaagtttttgttgaataatagtttcaaaattcaaaaacaataaatatgctcaatttagagcgtagagtaattttctcatgacctaataattctgcagtttagatttaatcatagtatctcatgatttgataattcaaaaatcgaaattaatacatactcagttgatgcttgtacattcacacgggttaatttctaccggtgtccaagcggtgtagcaagatcaaatagtatgggttagactcaatttacactttgtagtcaaagttgtgtgaccttgtaagcaatctagacaaatggcaaatctatggctatctaaaatttaaaacaaatcagacatgctatttacgggaagctgttttgatccctcaagagaatgtccctcgcttgtaaaaaaaaaccatctagaattctcaaaaaaaattcacaacattcatactatgtatgtatgtatgtatgcatgcatgcatgcatgcacacacacacacatgtatgtacatataacacatatacatatacatatacatattcatacataacgcacacacacaaattaacttaaatttttgaaaattttcaacttatgagtatatactaaggtacaagaatttgattcgtacctatatctagcaacaaaacaactcaaattcaactttattttacaatccattattacatacctaactaattatatgtttgggtggtatatacctagaagcaaaatctaacaaaaaaatgttcaaatttagttcaaacttgttagtaaagaagttaatgtccatacctgatctacattgccttgctctcatcatcgtcagcgttgtggcctttctcacaggcaacctctcctcgctcgagatccataaaaacaagagagaaaaaaatggattggttagttcgtgtgtgtatctatatataggtctatagtaaaaaaattactgggtgatgcgttgtgggatgggcgatgctggtagtctgcgggggtgggtttacacacacaacactccactaggtccaaactcaatttataacatttaggtataaaaaataaatatattcataccggaatttgtctttttgtttctcaatatataggtcgaatttgaagttgttctctcgtggaatgttagatgtcactgtactccacattgttaattaaatttttataactattttaattgaatttaaaagcaaaaaagtatatgaggggatatcctttcgtgggattagaattcattcctgctattaactagacctgtattgcagcgtgtttcctgaagattatactatgtcaagggagcaacgggtcgcagaaggcttcgtagtgggcattgaaaagtgcagggcagaggatgcggccgagcgaataatcaggaatgtgcctgtggactatgatgggcttggccaggtgagcacatccaagatacaggaccacttgcctgaacttgctcctcgtgctgtgcaaaacgagaagttcggctcttcaaatgatcttagcataatgatacaaatcaacaagtatgctcgtctgccatcttatgagtggagggacactgatatatccaaactgaatttccggcttctccgcgccccgatgttacttgaagctgtaacggcccgttgtcacttgttggacttaatcctcattggttcaaacaacattactgttcttgaccttggacgtcccacaatcaccaaactgccggcatctatagaatgtctacctaatttacggtacttgaggttgcagggcacgcaactgaagtcgctgtcggaggttattgtgaagatgcccaccatcagggggttggacatcaagaataccaagacagaggagctaccacaaggcattttgaggatgaagaagttgagtcacctgagtatgggggagaagcagaaaaatattcaagtattcatggaaaaaatgcagactctggctgagaccgtccaagacagcgatgacttgtccgacgaaacagagggaatagccgacgacgaaggcgagttctctacaagggccaacgcgtcaacaccgaaggtggatgaggacgaggtggaccggagggccaataacttcatcgccaggttcaggaagcagatcacgatcaggaattcaggattcgccaaaaaggagagcagcatcgacgaacgactgtggatacgagatctcgatgagtgccaactctccaagagaggtggacgattttaagaagaagttcgacgagatcattgccaggaacaggcacacatggaagccaatcgagtcaccgcgttctgtcaagcacggcaagtatttcgttcgctgcccgccgtcgctaatcaccagcgctgtcttttcctttttctttctttctatttttctagtatggtactccctccgtttcaggatattgtcaggaaacccctaatcaatcgccttgtgaacactggatgaactcacgcgaaagtggaagaacacgtagaacacaatagtgtagacgaacagacaacaaaaacagagtttttggtgctgcctgaattgcttgcagctttt</dnaseqindica> 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