Difference between revisions of "Os01g0136400"
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===Evolution=== | ===Evolution=== | ||
| − | The wall-associated kinase (WAK) gene family, one of the receptor-like kinase (RLK) gene families in plants, plays important roles in cell expansion, pathogen resistance, and heavy-metal stress tolerance in Arabidopsis (Arabidopsis thaliana). Through a reiterative database search and manual reannotation, we identified 125 OsWAK gene family members from rice (Oryza sativa) japonica cv Nipponbare; 37 (approximately 30%) OsWAKs were corrected/reannotated from earlier automated annotations. Of the 125 OsWAKs, 67 are receptor-like kinases, 28 receptor-like cytoplasmic kinases, 13 receptor-like proteins, 12 short genes, and five pseudogenes. The two-intron gene structure of the Arabidopsis WAK/WAK-Likes is generally conserved in OsWAKs; however, extra/missed introns were observed in some OsWAKs either in extracellular regions or in protein kinase domains. In addition to the 38 OsWAKs with full-length cDNA sequences and the 11 with rice expressed sequence tag sequences, gene expression analyses, using tiling-microarray analysis of the 20 OsWAKs on chromosome 10 and reverse transcription-PCR analysis for five OsWAKs, indicate that the majority of identified OsWAKs are likely expressed in rice.[4] Phylogenetic analyses of OsWAKs, Arabidopsis WAK/WAK-Likes, and barley (Hordeum vulgare) HvWAKs show that the OsWAK gene family expanded in the rice genome due to lineage-specific expansion of the family in monocots. Localized gene duplications appear to be the primary genetic event in OsWAK gene family expansion and the 125 OsWAKs, present on all 12 chromosomes, are mostly clustered. | + | The wall-associated kinase (WAK) gene family, one of the receptor-like kinase (RLK) gene families in plants, plays important roles in cell expansion, pathogen resistance, and heavy-metal stress tolerance in Arabidopsis (Arabidopsis thaliana). Through a reiterative database search and manual reannotation, we identified 125 OsWAK gene family members from rice (Oryza sativa) japonica cv Nipponbare; 37 (approximately 30%) OsWAKs were corrected/reannotated from earlier automated annotations. Of the 125 OsWAKs, 67 are receptor-like kinases, 28 receptor-like cytoplasmic kinases, 13 receptor-like proteins, 12 short genes, and five pseudogenes.[[File:table 1.jpg]] The two-intron gene structure of the Arabidopsis WAK/WAK-Likes is generally conserved in OsWAKs; however, extra/missed introns were observed in some OsWAKs either in extracellular regions or in protein kinase domains. In addition to the 38 OsWAKs with full-length cDNA sequences and the 11 with rice expressed sequence tag sequences, gene expression analyses, using tiling-microarray analysis of the 20 OsWAKs on chromosome 10 and reverse transcription-PCR analysis for five OsWAKs, indicate that the majority of identified OsWAKs are likely expressed in rice.[4] Phylogenetic analyses of OsWAKs, Arabidopsis WAK/WAK-Likes, and barley (Hordeum vulgare) HvWAKs show that the OsWAK gene family expanded in the rice genome due to lineage-specific expansion of the family in monocots. Localized gene duplications appear to be the primary genetic event in OsWAK gene family expansion and the 125 OsWAKs, present on all 12 chromosomes, are mostly clustered. |
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==Labs working on this gene== | ==Labs working on this gene== | ||
Revision as of 12:48, 28 May 2014
The expression product of Os01g0136400 is wall-associated kinase 1 (WAK1) which is a transmembrane protein containing a cytoplasmic Ser/Thr kinase domain and an extracellular domain in contact with the pectin fraction of the plant cell walls.
Contents
Annotated Information
Function
plays important roles in rice blast disease resistance
Wall-associated protein kinases (WAKs) can phosphorylate OsRFP1, a putative transcription regulator recently identified in rice. OsRFP1 strongly interacts with the kinase domain of OsWAK1. This demonstrated that OsWAK1 is a functional protein kinase. A fusion protein of OsWAK1 with GFP was found to be localized on the cell surface. Northern blotting analysis showed that infection of the rice blast fungus, Magnaporthe oryzae significantly induced the OsWAK1 transcripts, and the accumulation of OsWAK1 mRNA occurred earlier and was more abundant in rice leaves infected with an incompatible race than with a compatible race of the blast fungus. OsWAK1 was also induced after treatment by mechanical wounding, SA and MeJA, but not by ABA. These results imply that OsWAK1 is a gene involved in plant defense. Furthermore, six transgenic rice lines with constitutive expression of OsWAK1 became resistant to the compatible race. However, OsWAK1 expression was undetectable in leaves, stems and flowers but very weak in roots under normal growth conditions. This provides functional evidence that induction of OsWAK1 as novel RLK plays important roles in plant disease resistance.[1] Oligogalacturonides (OGs) released from the plant cell wall are active both as damage-associated molecular patterns (DAMPs) for the activation of the plant immune response and regulators of plant growth and development. Members of the Wall-Associated Kinase (WAK) family are candidate receptors of OGs, due to their ability to bind in vitro these oligosaccharides. Because lethality and redundancy have hampered the study of WAKs by reverse genetics, we have adopted a chimeric receptor approach to elucidate the role of Arabidopsis WAK1.[2]
Expression
A comparative analysis on protein kinases encoded in the completely sequenced genomes of two plant species, namely Arabidopsis thaliana and Oryza sativa spp japonica cv. Nipponbare is reported in the current study. We have analysed 836 and 1386 kinases identified from A. thaliana and the O. sativa genomes respectively. Their classification into known subfamilies reveals selective expansions of the plant receptor kinase subfamily comprising of Ser/Thr receptor kinases. The presence of calcium dependent kinases, and potential absence of cyclic nucleotide-dependent protein kinase of the type found in other (non-plant) eukaryotes, are other notable features of the two plant kinomes described here.
An analysis on domain organisation of each of the protein kinases encoded in the plant genome has been carried out. Uncommon composition of functional domains like nuclear translocation factor domain, redox sensor domain (PAS), ACT and lectin domains are observed in few protein kinases shared between the two plant species. Biochemical functions characteristic of the domains recruited in these protein kinase gene products suggest their mode of regulation by alternate cellular localisation, oxidation potential, amino acid flux and binding of carbohydrates. Occurrence of multi-functional kinases with diverse enzymatic modules, such as Transposases and peptidases, tethered to the kinase catalytic domain is another interesting feature of the protein kinase complement of the O. sativa genome. Co-occurrence of diverse nucleotide and carbohydrate binding domains with catalytic kinase domain containing gene products has also been observed. Putative homologues of protein kinases of A. thaliana that regulate plant-specific physiological processes like ethylene hormone response, somatic embryogenesis and pathogen defence have been identified in O. sativa genome as well.[3] Intron-exon structure is conserved between theWAK and WAKL genes. The diagram shows a standardized depiction of a WAK or WAKL gene from each of the four groups (I-IV). Exons are represented by boxes. Introns are represented as ‘V’s. Regions of each gene encoding functional domains are indicated with shaded boxes: N-terminal signal sequence (black), EGF2-like domain (red), calcium-binding EGF domain (blue), transmembrane domain (green), and Ser/Thr protein kinase active site (orange).
Evolution
The wall-associated kinase (WAK) gene family, one of the receptor-like kinase (RLK) gene families in plants, plays important roles in cell expansion, pathogen resistance, and heavy-metal stress tolerance in Arabidopsis (Arabidopsis thaliana). Through a reiterative database search and manual reannotation, we identified 125 OsWAK gene family members from rice (Oryza sativa) japonica cv Nipponbare; 37 (approximately 30%) OsWAKs were corrected/reannotated from earlier automated annotations. Of the 125 OsWAKs, 67 are receptor-like kinases, 28 receptor-like cytoplasmic kinases, 13 receptor-like proteins, 12 short genes, and five pseudogenes.
The two-intron gene structure of the Arabidopsis WAK/WAK-Likes is generally conserved in OsWAKs; however, extra/missed introns were observed in some OsWAKs either in extracellular regions or in protein kinase domains. In addition to the 38 OsWAKs with full-length cDNA sequences and the 11 with rice expressed sequence tag sequences, gene expression analyses, using tiling-microarray analysis of the 20 OsWAKs on chromosome 10 and reverse transcription-PCR analysis for five OsWAKs, indicate that the majority of identified OsWAKs are likely expressed in rice.[4] Phylogenetic analyses of OsWAKs, Arabidopsis WAK/WAK-Likes, and barley (Hordeum vulgare) HvWAKs show that the OsWAK gene family expanded in the rice genome due to lineage-specific expansion of the family in monocots. Localized gene duplications appear to be the primary genetic event in OsWAK gene family expansion and the 125 OsWAKs, present on all 12 chromosomes, are mostly clustered.
Labs working on this gene
1.Department of Horticulture, Michigan State University, East Lansing, Michigan 48824
2.Department of Biology, San Francisco State University, San Francisco, California 94132
3.Department of Plant and Microbial Biology, University of California, Berkeley, California 94720
4.Department of Molecular, Cellular, and Developmental Biology, Yale University, New Haven, Connecticut 06520–8104
References
1.Santy Peraza-Echeverria, Andrew James-Kay, Blondy Canto-Canché, Eduardo Castillo-Castro (2007).Structural and phylogenetic analysis of Pto-type disease resistance gene candidates in banana. Molecular Genetics and Genomics 278,443-453.
2.Alexandre Brutusa; Francesca Siciliaa, Alberto Maconeb, Felice Cervonea, and Giulia De Lorenzoa (2010). A domain swap approach reveals a role of the plant wall-associated kinase 1 (WAK1) as a receptor of oligogalacturonides. Alexandre Brutus, doi: 10.1073.
3.A. Krupa, Anamika, and N. Srinivasan (2006).Genome-wide comparative analyses of domain organisation of repertoires of protein kinases of Arabidopsis thaliana and Oryza sativa. elsevier 380,1-13.
4.Shibo Zhang, Calvin Chen, Lei Li, Ling Meng, Jaswinder Singh, Ning Jiang, Xingwang Deng, Zhenghui He and Peggy G. Lemaux (2005). Evolutionary Expansion, Gene Structure, and Expression of the Rice Wall-Associated Kinase Gene Family. American Society of Plant Biologists 139,07-24.
Structured Information
| Gene Name |
Os01g0136400 |
|---|---|
| Description |
Protein kinase-like domain containing protein |
| Version |
NM_001048492.1 GI:115434397 GeneID:4325700 |
| Length |
9605 bp |
| Definition |
Oryza sativa Japonica Group Os01g0136400, 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 1:1956847..1966451 |
| Sequence Coding Region |
1957104..1958041,1958213..1958467,1958587..1958619,1965545..1966325 |
| Expression | |
| Genome Context |
<gbrowseImage1> name=NC_008394:1956847..1966451 source=RiceChromosome01 preset=GeneLocation </gbrowseImage1> |
| Gene Structure |
<gbrowseImage2> name=NC_008394:1956847..1966451 source=RiceChromosome01 preset=GeneLocation </gbrowseImage2> |
| Coding Sequence |
<cdnaseq>atgcacccgaccttgttgtgcttgccactcttggcctccttgctcctcctgtgccaccgcgcgcgcgccgagtgcgagccggcgacatgcggcaacctcaccgtcaggtacccattctggctaggcggccccaacttcaaccagtccaaccagtcgtcgccgtcgtcagctctggcctcctgtggccatccggctttcgaggtgtggtgtaacggcggcgtggcgtcgttgaggggctcgcaaatcctcgtcctcagcatcgactacaacagcagctcattcgtcgcggcacacaaaagggtcgccgacggcggcgacggcgtgtgccgcaccgacttcaacatatcgtccagcctagccctcagcccgttcacgatcagcagcagcaaccgggccatctgcttcctctacagctgcaacggcacggagccaccggagatcgacggccttgtgaacgccaccatctccagctgcagtaagcctatctacgcgtacctcggcgggatctacgaccgtgataatccaccggcgatcaaagccgggaactgcacgtactcgtacctgccggtgctgtggccggactcgccggcgaacttgacggcggggacaaactacagcccgcagttcaagaaggggttcgtgttggagtggcagaagaacgggttcggcgactgcgacgcctgtaacgggagcggcggtcagtgccggtacatcaacgattccgcggcggcgttcgcgtgcctctgctccgacggcaagctgcgccgctcgacatgccccggttcaagaagcaagagtcacataataggaatagcatgtggatcaagcggcggaatattattgattgtatctatattcatttttgcttggcacaaacgcaagaagaggaaacaaacccgagatttgaaagatctcatgcatagttcatcttcaatgcaatcatacagcaaagaccttgagttgggtggttctccccatatattcacttacgaggaacttgaagaggctactgctggatttagtgcctcgagggaacttggtgatggtggttttggaactgtttacaaaggaaagctccgggatgggagagtagttgcagtgaagcgcctttacaagaacaactacagacgagtagagcaattcctaaatgaggtagacattttgtcccgcctactgcaccagaaccttgttatcctatatggctgcacgtctcgttctagccgtgaccttctcttggtctatgagtacatcccaaatgggacagttgcagaccatctacatggaccccgtgcaggagaacgaggcctcacatggcctgtaagaatgacaattgcgatagaaacggctgaggcactggcataccttcatgcagttgaaatcatacaccgtgatgtcaagaccaacaacatattgctggacaacaacttccatgtcaaagttgcggactttggactatcgcgcctgttcccgcttgaagtcacccatgtatcaactgttccacagggcacaccagggtatgttgacccagtgtaccaccagtgctacaagctaaccgataagagtgatgtgtatagctttggtgttgtgttgatagagctaatttcctcaaaaccagctgtggacatgtccaggagccacagtgacattaacttggctaacatggctctcaacagaattcagaaccatgaagttgatcagttggttgatccagagatcggctatgagactgacagtgaaacaaagaggatggtagatctggtggccgagctggcctttcagtgcttgcagatggacagagagagcaggccaccaattaaggaggtagtggaggtcctgaattgtatcaagaacggggaatgtccagcggaaaagatgaacaagaatgcgtctccaaaggaagattcgcatctgctgaaggacagcctacagtattcgcctgactcagtaatccatagatttcatagccaatctactaaccactcggtagcatcaaactctagcggatga</cdnaseq> |
| Protein Sequence |
<aaseq>MHPTLLCLPLLASLLLLCHRARAECEPATCGNLTVRYPFWLGGP NFNQSNQSSPSSALASCGHPAFEVWCNGGVASLRGSQILVLSIDYNSSSFVAAHKRVA DGGDGVCRTDFNISSSLALSPFTISSSNRAICFLYSCNGTEPPEIDGLVNATISSCSK PIYAYLGGIYDRDNPPAIKAGNCTYSYLPVLWPDSPANLTAGTNYSPQFKKGFVLEWQ KNGFGDCDACNGSGGQCRYINDSAAAFACLCSDGKLRRSTCPGSRSKSHIIGIACGSS GGILLIVSIFIFAWHKRKKRKQTRDLKDLMHSSSSMQSYSKDLELGGSPHIFTYEELE EATAGFSASRELGDGGFGTVYKGKLRDGRVVAVKRLYKNNYRRVEQFLNEVDILSRLL HQNLVILYGCTSRSSRDLLLVYEYIPNGTVADHLHGPRAGERGLTWPVRMTIAIETAE ALAYLHAVEIIHRDVKTNNILLDNNFHVKVADFGLSRLFPLEVTHVSTVPQGTPGYVD PVYHQCYKLTDKSDVYSFGVVLIELISSKPAVDMSRSHSDINLANMALNRIQNHEVDQ LVDPEIGYETDSETKRMVDLVAELAFQCLQMDRESRPPIKEVVEVLNCIKNGECPAEK MNKNASPKEDSHLLKDSLQYSPDSVIHRFHSQSTNHSVASNSSG</aaseq> |
| Gene Sequence |
<dnaseqindica>8411..9348#7985..8239#7833..7865#127..907#agtcaagaaaaacaaggcaatggcaatagcatttctcgagcattagaccttcccctccctttccagcattctgctgctctccaatccggcctccattgtgtagcagctagctccacgagcggcaagatgcacccgaccttgttgtgcttgccactcttggcctccttgctcctcctgtgccaccgcgcgcgcgccgagtgcgagccggcgacatgcggcaacctcaccgtcaggtacccattctggctaggcggccccaacttcaaccagtccaaccagtcgtcgccgtcgtcagctctggcctcctgtggccatccggctttcgaggtgtggtgtaacggcggcgtggcgtcgttgaggggctcgcaaatcctcgtcctcagcatcgactacaacagcagctcattcgtcgcggcacacaaaagggtcgccgacggcggcgacggcgtgtgccgcaccgacttcaacatatcgtccagcctagccctcagcccgttcacgatcagcagcagcaaccgggccatctgcttcctctacagctgcaacggcacggagccaccggagatcgacggccttgtgaacgccaccatctccagctgcagtaagcctatctacgcgtacctcggcgggatctacgaccgtgataatccaccggcgatcaaagccgggaactgcacgtactcgtacctgccggtgctgtggccggactcgccggcgaacttgacggcggggacaaactacagcccgcagttcaagaaggggttcgtgttggagtggcagaagaacgggttcggcgactgcgacgcctgtaacgggagcggcggtcagtgccggtacatcaacgattccgcggcggcgttcgcgtgcctctgctccgacggcaagctgcgccgctcgacatgccccggtgagtaccacacttgcctatcgcccaatctgacgcatgtgttagctagttgaagattttggctcggtgtacggttaactggacggccaaattggcgcgttgcgatcggtagccatgtactccgatcccaaagttgcaatctctcgtaggtcccatgtggatttggagttgaattactggcccacgtcaatgtccttggattttactcaacctctcgctcgctcgcggtctcgggtagttacatgggcggctggctagtcagctgcagagatggtggtaaattttgggagggtttgcctcggtggtgacgttgacgacgacttatcaagcaggctgcaggcaggagttgacatgggcggctagtcactggttcttccattgaccacccagcgtgattgattcttgggtaaacgcatgaattccgtcctttcctttgtctctgtctcccaacttcccagggcagagatccttctgatccttctgcttccccctcttctttttcccccgaaaattcaccattctcattagtgaactggttcattattaggacaaataagacgttagcagtagttgcctgaagcttcgagtgtgttgagctggtccataaactaatgctcatagtaggtgacaatgtaaatcattagtacatgatcaccaagtggcattctgcgtagaagtaccaaaatagtggggacagatgggcacgggctaaatttggcatcgtgcacggcgcaccagccgctaccaggggctttctttgaactgataaggctaacgggataagttacgctttatatctttcgaaagcggtcaatcggagttgttagaagtggcgggccatgaatttttggagccgaaagaagcgaatattaaggaaagacctaatatcaaataattagaagggttgatactttgaacccagatcgtctcgttcaccatcttgtggagctagccggaagaccctggacgtttctcaaatttttggagcccaacatatacgcagacataattggaaacacaatggaaagctcaacatataagacatcataatggtgcgatctattggcttcttaaccttagttactgctccctccggtacaaaaatcttgatgttttaaccttgtaaatggtgattgcgaattgttaatgcttctatcttagcatgttatattggaccgctttgccctacatgcttggaagtaaaaaccaagtaaaaaagctagggaattggacctttagctcgcaccgatacaggtgattaattcaaaccaacggcagcagtgtagtgattaccatgagtacaatactatatttaatatggtaaaatagtaaattgatactgtaatatggagtaaatcacttcttggtatgtgagatcatgttcaaaacatcaataattttgaaactggaaggagtacacactgcccttctcaacttttaaacgctactatttccaaaaagaaaactttaaaatgcaataaacagatatgaaaaactgttccctcaaaaaatatatgaaaaattcaaaacaggaaatgcggtagtgctagaaatatgaaattatgaactttaatcagtgtagtttgtattgaaattttaatacaatggggaatgaaagcttatataaaaataaaaaatatatagaaaatgcaaaacaatatactaccttggtgcccccattaacgatagtactagagattgccaccactaagctaattccggtctatgagtctatccaagtgcttttacataaaagagcattgtgagctttgctgataatgacatagatttcggcaaaatatggtaccaaatgctaatgcctttcctgcaaagattctgattctaggagttgatagatacctacataatttttaggcttacttttttttcaaaattactttccaaattgttgaaatgaattttgtgctctacgctcctaaaatatattttctcaaaaaaatattctattgtattgcacacttgttttaagttcatttattaaatgtttaatttataactacaatcaagtagataatctgtacaacaatccacttaataatccttagattgtgtttgagtgatgaaatgagaaaattaagaagatacaacaaacaagatgagtcattagcttatatgaattgagtattaactgttttaaatttgaaactatatgaattgagtattactttattatatttatctataaacatagttaaacatgagacagtttgactgaccaaaaatcaaaacaacttataatctaaaacggagggagtactggttagtggtaggaactagcaacgcagcacatatgtgttcaatctgcattaggtttttggttagagagaagatactagatattaacacgtttttatcagaaagtactccttctgtcctatattattaggttcagaaaaatgtatctacaagttctcataatattaggatatgtcaaattaggtaggtttttattaaacacatgaagtattctgtaggagtttttttttagttttagttacatccgtacaggtaagcgttgaccggttggaaagtggactagtctctgcttaccggcaagagtaggtgtggaacaaagggcgtagtagcctgttatacaacaggtagaaaacaatggtggacagcagcgtgcactagctagctggaagagggttaataggagtacacgacgatgagtccggacaaactggtccagtcacatcagggcggttataacagctagctgcccatagggcaccccggaaactcgcacgcgtggacgacgatcgttcgtgtatcaggtcaaacatttcgacacagaggaatagttcccgtgtgcaaagtttcttacttccatctcccccacaaacttccatctccttcagaaatcccctatccccctcctcctctttctgtctcgatgtccccgagcttcttctttgtcgtcgtctcggcctggtcgctagcgctgatgctcgccgcggcggcgaggggagccgaggaggaaggaggaggaggctgcctgggcagccagaaatgcggcgacctgaatatctcctctccgttctggatcatccagggccaggcggataagccgtgtggtcctctggattaccaggtatattgcaacaactccaccggcgtcgcaactcttcgaagctctacagacagcgggtttgatatcatcaacatatcatatggggaccgtactatgctcgtctttgatgtccataagctagctcgcctgaataactccaccggctgcagtatcccagtgtttaacaccttcgccaagctgcccatcacgtttacaatcagcccttccaatcacaacctcgtcttctacaactgcaccgaggcgccgccggcggagcagcagcaacaactggggctcgtggagacgagatgcggtaacaacacgtttgctcgcctgggagggcgtttccacggggagggcgactacgacaagtactatttggaaggctgcagcagaaacagcaccgtcttcttgccggtgctggaaccgcctgatggcaaggcgaacgccagcaggtatgtggagctcgtgggtggaggcttcctcataacatgggacctgccaccgccagtgacatcttctggtaagttcaccctccctgaaactattaggatcaagttcgtatagaaaatccactgtatatcctgatacttccgatctccaagcgagtactagtagaatacggttctcgctcatcagcgtgtaggacagggaatctgtcgattggctgatagggtcgtcatcttgcccaatcgtggcggctgggcaacagggatgagcaagaattaaactagaattaaagtgatcaaagcagtggaccatgaccgtcctttcgcattccatctcactccggtgccattccttccaatcttcgatcctgttctggtacatgactgagaagaccatgcacatctgttggaattggaaccgatacggcaagacgatgaggccgtagacagccattatccagtcaattttttcggccactacagagaattcccaaagttaaatgtatcgtactagaagaagaagtagcagtattaattctctcatgtcttcctcaatcctcatcaacaacacatcaccaattcatcctcttcgttttcgatgccctctccttccttgttcctcttgttcgcctgcctcgcctgggcgagtcaagcagcgaatacggcggcagacaatcgtccacaagaaggctgcgcggccagtactgtatgtggcaaggtgaccatctcgtcgccgttcgccgtcgtgccggagcaggcaacggagagcaaatgcggctggcttggattccaggttatctgccacaacgacactccatacctcggctactacaagcccagatatcggatccagatcctcgacatcttctacggcaacaattcattgctcgtctctgacatccacaagctcggtgacttcattgtcttctccggcgtcagcaaagaatactcctgccatgttccgaggaccaacacctcctccaaggtcggcctcccgttctccatcagcaccaccaatctcaacctcttcctgtacagttgcaataaggcgcttgtgccgcgggacggagacgacgacctcgtggagacgaggtgcggcaacaagacgtttgctcgcgtaggagggaattacagtgattcgggcgactacccggcgttttacatggaaggctgcaatgctaccgtcgtgccggtgctgggcacggacgcgaggagctatgagcagctcatccgcgacggcttcctcttgacatggcaagagacgccgtcatctggtaagttcgttcgcgaaattatccatttaatcatcactttcgggaggaggaaatgcgtaaaatttatggtgtcaactttatcgaatccgttaatcaatcagtgacttgtttattggttgataaggcagtcgtcatcgctaatccatctccgacctttggatatcgaaatcagagaaatatatgcaaaaactatactcccttcgtttcacgatgtaaatcattttagcgtttcttacgttaatattgatgttaattaatctagacatatatacctatctagatttattaacatcaatataaacgtgaaaaatactatagtgatgaaacggagaaaatagtacggagtacttcgtcaagaagctgtagcagcagagagagtagtagcaatgtagcagtatgttcaatccgtagactttgaccggtcaaaaccgcttcgccaccgtccttttgccccacgcctcatatcccatccctcgatctctcgtccccttcttctctctcatataatccatttcaccactgcattccgtagttgatttgcgtacccaagtccaccagatatgcctccgctcatactgctactgctggtagcttccttcctcgagttgccggcaccggcgagctcgtctagtcctggctgcttgcccacgccatgcggcaagctgaccatctcctacccgttctggctggaggagcccggccggccgccgtgcgggtcgccgcccttccagctcaagtgcaacgccaccggcgcatacctcacgcacaccatctacgaggcgtatcgcgttgttgatatcttcaccggaaaccacaccgtccatgtggtggacgagaatctcccgctcgccaccggctgcccggcgccgccgttcaacatctccgatggcatctggcaggcgccgttcgtcatcagcgaagccaacgcagagctgcgcttcctctcgtgtaacaagtcacttccggcggcggctgctcctcccggcttccatagcctgccttgtgatgaccaaaactcctccgtccggctcgtcagcgaccaccatttacacgaggatgggattccaccgggctgtaacttcacggttgtgccgatcgttcagcgtcacaatgggagtatggccggctatattgccagcatgaggagtgggtttctactagagtgggcggtggtttcaggggattgtcccaaatgtcaagtaagcggcgggaattgcacgtacagcgacgacctggagttcgcctgcaattgccccgacgggatgcaccctgacaagtgtagagagttcagaaaatcggaagagcacggtaaatttgtccagtcaattatcaattagaataaaaattttaacatagttctgtgaacatgcagcctaggccatcctaaccgattcaatgcatccaaagtgtgttcatctagttgatagttttctttatatatatagaaaagaatgacacatcatttcgttatacaggaagatttccatgctactgtggcctaatgtccactagataaactcttattataataatgcgcgtgacatctagttttgctatcgaggaaagaaattaaacacagcattgtcacagcaatgttcataagaatataaaatgcaaagtggatattgatatttttgttttgaattatgagtgtttgcttcaagttcctgctgttttggttgttagtttctcatcaaacaattatttcactgatgggaacattggcatttggcagcttatggaatcctagtcaaacgtgttacgtgtagtttgaagtattcaactacactatctgataaagatatcataccaataccatgttccaatttttcttctgctgattgctgactgacgaagaaatccctcatctgattgacagcaaataccctttcattttctaatccttaattagtgtgttccaaacatgtggaaagcataaattgcatctgtatttctgttgaattaggctgaaaagttgtataagcactgtccttcgtttcaacagggggttaggggagcaatgataccctaacaatgtgattatgaatgatcattctatttgttcatggcatatttgtgcaaggtattaaaagtctgttcttctgcaggttcaagaagcaagagtcacataataggaataggtgagtataattcattcttgtttcttggacttccccaaccattcaaaatgtaattatgaagatatgcttcagatccttggttctcacatattgatagtgctctgtctctgttattgcagcatgtggatcaagcggcggaatattattgattgtatctatattcatttttgcttggcacaaacgcaagaagaggaaacaaacccgagatttgaaagatctcatgcatagttcatcttcaatgcaatcatacagcaaagaccttgagttgggtggttctccccatatattcacttacgaggaacttgaagaggctactgctggatttagtgcctcgagggaacttggtgatggtggttttggaactgtttacaaaggtaggagaatatgtttgcacaactcccttgttgttaatagatttctcatctaatcacttcctcattagcatgcctaattgcatgcgacaggggctccgttgaactttgttccataatagataacttcccctattaacttagcagttacaattcttcctttctgccattcaggaaagctccgggatgggagagtagttgcagtgaagcgcctttacaagaacaactacagacgagtagagcaattcctaaatgaggtagacattttgtcccgcctactgcaccagaaccttgttatcctatatggctgcacgtctcgttctagccgtgaccttctcttggtctatgagtacatcccaaatgggacagttgcagaccatctacatggaccccgtgcaggagaacgaggcctcacatggcctgtaagaatgacaattgcgatagaaacggctgaggcactggcataccttcatgcagttgaaatcatacaccgtgatgtcaagaccaacaacatattgctggacaacaacttccatgtcaaagttgcggactttggactatcgcgcctgttcccgcttgaagtcacccatgtatcaactgttccacagggcacaccagggtatgttgacccagtgtaccaccagtgctacaagctaaccgataagagtgatgtgtatagctttggtgttgtgttgatagagctaatttcctcaaaaccagctgtggacatgtccaggagccacagtgacattaacttggctaacatggctctcaacagaattcagaaccatgaagttgatcagttggttgatccagagatcggctatgagactgacagtgaaacaaagaggatggtagatctggtggccgagctggcctttcagtgcttgcagatggacagagagagcaggccaccaattaaggaggtagtggaggtcctgaattgtatcaagaacggggaatgtccagcggaaaagatgaacaagaatgcgtctccaaaggaagattcgcatctgctgaaggacagcctacagtattcgcctgactcagtaatccatagatttcatagccaatctactaaccactcggtagcatcaaactctagcggatgatgagaaactttgtattgattctgatgaaataaggataactaagttgtcccctcttggttgatggactaattaatactacaagtgtttgactatgagcttggcagaaattcagacaattggtgaatgcggtaggagaaatacagggttcaaacaatctcgtgtaatctagtgttcagtgtgttcatgatttcaggttagcgagtggtatcagggagaacacatcccattgttaccctaactagaggttgcaattttgc</dnaseqindica> 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