Difference between revisions of "Os09g0401100"
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exon by 819 nt (Figure 1a). The first exon of antiSTR1 matches 131 nt of another gene adjacent to STR1 (RAP ID, Os09t0401200-01; data not shown) in sense orientation. AntiSTR1 is predicted (http://www.rapdb.dna.affrc.go.jp) to encode a protein of 94 amino acids. However, no corre- sponding protein sequence has been reported from any organism, as determined by BLAST searches. Therefore, antiSTR1 is not likely to encode for a protein but instead represents a bona fide cis-NAT. Computational searches for similar cis-NAT sequences from other plant species did not yield any match. The presence of antiSTR1 was confirmed by real-time RT-PCR, with primers specifically targeting antiSTR1 and with cDNA synthesized by an antiSTR1- specific primer (Tables S1 and S3). Natural antisense transcripts have been proposed to regulate complementary mRNA in many different ways, such as transcriptional interference, RNA masking, chro- matin remodelling, RNA editing and RNA interference (Lapidot and Pilpel, 2006). For RNA interference, small interfering (si) RNAs of mostly 21–25 nt in length are produced from the antisense transcript, and lead to Argo- naut-dependent degradation of mRNAs (Okamura and Lai, 2008). We considered the possibility that siRNAs might be produced from antiSTR1 and could also target other genes in trans. Applying a cut-off of 21-nt sequence complemen- tarity revealed a matching sequence of 121 nt in length contained within the first exon of the STR2 open reading frame, indicating the possibility that STR2 could be targeted by antiSTR1 in trans. However, as antiSTR1 originates from the same locus as STR1 we kept the name antiSTR1. Small complementary stretches of 23–34 nucleotides were also found in three other genes located on chromosomes 4 and 7, and encoding an inorganic pyrophosphatase, a glycosyl- transferase and a hypothetical protein (Table S2). However, these genes are not regulated by AM colonization (Gu ¨imil et al., 2005; C. Gutjahr, R. J. H. Sawers, H. Angliker, T. Roloff, E. Oakeley, U. Paszkowski unpublished data). Next, to confirm the AM-responsive induction of STR1, and to assess AM induction of STR2 and antiSTR1, real-time RT-PCR analysis was performed on cDNA from non-colo- nized and Glomus intraradices-colonized roots. High levels of STR1 and STR2 mRNA were detected in mycorrhizal roots, thereby confirming previous microarray data, whereas expression was at the background level in non- colonized roots (Gu ¨imil et al., 2005; C. Gutjahr, R. J. H. Sawers, H. Angliker, T. Roloff, E. Oakekly, U. Paszkowski, unpublished data). antiSTR1 mRNA also accumulated in roots, but the level did not change upon AM colonization (Figure 1b). Examination of the transcript levels of STR1, STR2 and antiSTR1 in panicle, leaf, stem and embryo revealed only background levels of expression in these organs for all three genes. As STR1 and STR2 were induced in mycorrhizal roots we next examined their expression in arbusculated cells collected by laser microdissection (Figure 1c). Transcript levels were compared between arbusculated and non- arbusculated cortex cells from mycorrhizal roots (hereafter called systemic cortex cells) and cortex cells from mock- inoculated roots. To ensure the specificity of the samples, we first recorded the expression of the arbuscule marker PT11. PT11 was specifically expressed in arbusculated cells, but not in systemic or mock control cells, as reported earlier (Gutjahr et al., 2008). Similarly, STR1 and STR2 were | exon by 819 nt (Figure 1a). The first exon of antiSTR1 matches 131 nt of another gene adjacent to STR1 (RAP ID, Os09t0401200-01; data not shown) in sense orientation. AntiSTR1 is predicted (http://www.rapdb.dna.affrc.go.jp) to encode a protein of 94 amino acids. However, no corre- sponding protein sequence has been reported from any organism, as determined by BLAST searches. Therefore, antiSTR1 is not likely to encode for a protein but instead represents a bona fide cis-NAT. Computational searches for similar cis-NAT sequences from other plant species did not yield any match. The presence of antiSTR1 was confirmed by real-time RT-PCR, with primers specifically targeting antiSTR1 and with cDNA synthesized by an antiSTR1- specific primer (Tables S1 and S3). Natural antisense transcripts have been proposed to regulate complementary mRNA in many different ways, such as transcriptional interference, RNA masking, chro- matin remodelling, RNA editing and RNA interference (Lapidot and Pilpel, 2006). For RNA interference, small interfering (si) RNAs of mostly 21–25 nt in length are produced from the antisense transcript, and lead to Argo- naut-dependent degradation of mRNAs (Okamura and Lai, 2008). We considered the possibility that siRNAs might be produced from antiSTR1 and could also target other genes in trans. Applying a cut-off of 21-nt sequence complemen- tarity revealed a matching sequence of 121 nt in length contained within the first exon of the STR2 open reading frame, indicating the possibility that STR2 could be targeted by antiSTR1 in trans. However, as antiSTR1 originates from the same locus as STR1 we kept the name antiSTR1. Small complementary stretches of 23–34 nucleotides were also found in three other genes located on chromosomes 4 and 7, and encoding an inorganic pyrophosphatase, a glycosyl- transferase and a hypothetical protein (Table S2). However, these genes are not regulated by AM colonization (Gu ¨imil et al., 2005; C. Gutjahr, R. J. H. Sawers, H. Angliker, T. Roloff, E. Oakeley, U. Paszkowski unpublished data). Next, to confirm the AM-responsive induction of STR1, and to assess AM induction of STR2 and antiSTR1, real-time RT-PCR analysis was performed on cDNA from non-colo- nized and Glomus intraradices-colonized roots. High levels of STR1 and STR2 mRNA were detected in mycorrhizal roots, thereby confirming previous microarray data, whereas expression was at the background level in non- colonized roots (Gu ¨imil et al., 2005; C. Gutjahr, R. J. H. Sawers, H. Angliker, T. Roloff, E. Oakekly, U. Paszkowski, unpublished data). antiSTR1 mRNA also accumulated in roots, but the level did not change upon AM colonization (Figure 1b). Examination of the transcript levels of STR1, STR2 and antiSTR1 in panicle, leaf, stem and embryo revealed only background levels of expression in these organs for all three genes. As STR1 and STR2 were induced in mycorrhizal roots we next examined their expression in arbusculated cells collected by laser microdissection (Figure 1c). Transcript levels were compared between arbusculated and non- arbusculated cortex cells from mycorrhizal roots (hereafter called systemic cortex cells) and cortex cells from mock- inoculated roots. To ensure the specificity of the samples, we first recorded the expression of the arbuscule marker PT11. PT11 was specifically expressed in arbusculated cells, but not in systemic or mock control cells, as reported earlier (Gutjahr et al., 2008). Similarly, STR1 and STR2 were | ||
specifically expressed in arbusculated cortex cells. Arbus- cule-specific expression was also observed for antiSTR1. This cell-specific expression pattern was unexpected as the expression level of antiSTR1 was similar between mock and Glomus intraradices-inoculated root systems (Figure 1b). antiSTR1 transcript accumulation might therefore differ spatially between colonized and non-colonized roots, but the total level of antiSTR1 transcripts in the root system might not differ between the two treatments. In summary, | specifically expressed in arbusculated cortex cells. Arbus- cule-specific expression was also observed for antiSTR1. This cell-specific expression pattern was unexpected as the expression level of antiSTR1 was similar between mock and Glomus intraradices-inoculated root systems (Figure 1b). antiSTR1 transcript accumulation might therefore differ spatially between colonized and non-colonized roots, but the total level of antiSTR1 transcripts in the root system might not differ between the two treatments. In summary, | ||
| − | all three transcripts accumulate in arbusculated cells, suggesting they play a role in arbuscule development, maintenance or function.[[File:1111. | + | all three transcripts accumulate in arbusculated cells, suggesting they play a role in arbuscule development, maintenance or function.[[File:1111.png|200px|thumb|left|alt text]] |
===Evolution=== | ===Evolution=== | ||
Revision as of 08:47, 25 May 2014
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Contents
Annotated Information
Function
Please input function information here. The half-size ABC transporters STR1 and STR2 are indis- pensable for mycorrhizal arbuscule formation in rice The central structureof thesymbiotic association betweenplants and arbuscularmycorrhizal (AM) fungi is the fungal arbuscule that delivers minerals to the plant. Our earlier transcriptome analyses identified two half-size ABCG transporters that displayed enhanced mRNA levels in mycorrhizal roots. We now show specific transcript accumulation in arbusculated cells of both genes during symbiosis. Presently, arbuscule-relevant factors from monocotyledons have not been reported. Mutation of either of the Oryza sativa (rice) ABCG transporters blocked arbuscule growth of different AM fungi at a small and stunted stage, recapitulating the phenotype of Medicago truncatula stunted arbuscule 1 and 2 (str1 and str2) mutants that are deficient in homologous ABCG genes. This phenotypic resemblance and phylogenetic analysis suggest functional conservation of STR1 and STR2 across the angiosperms. Malnutrition of the fungus underlying limited arbuscular growth was excluded by the absence of complementation of the str1 phenotype by wild-type nurse plants. Furthermore, plant AM signaling was found to be intact, as arbuscule-induced marker transcript accumulation was not affected in str1 mutants. Strigolactones have previously been hypothesized to operate as intracellular hyphal branching signals and possible substrates of STR1 and STR2. However, full arbuscule development in the strigolactone biosynthesis mutants d10 and d17 suggested strigolactones to be unlikely substrates of STR1/STR2. Interestingly, rice STR1 is associated with a cis-natural antisense transcript (antiSTR1). Analogous to STR1 and STR2, at the root cortex level, the antiSTR1 transcript is specifically detected in arbusculated cells, suggesting unexpected modes of STR1 regulation in rice.
Expression
Please input expression information here. Transcripts of STR1, STR2 and antiSTR1 accumulate in arbusculated cells [[File:To detect mycorrhiza-regulated genes with a possible func- tion in AM development in monocotyledons, we previously performed whole-genome microarray analysis of mycorrhi- zal rice roots (Gu ¨imil et al., 2005). One strongly AM-induced gene encoded an ABCG transporter homologous to Medi- cago STR1 (Zhang et al., 2010). Different versions of the STR1 transcript (TIGR ID, LOC_Os09g23640; RAP ID, Os09g0401100)werefoundinricegenomedatabases(http:// www.orygenesdb.cirad.fr, http://www.rice.plantbiology.msu. edu and http://www.rapdb.dna.affrc.go.jp). Sequencing of the cDNA from mycorrhizal roots revealed that the STR1 gene consists of four exons that correspond to LOC_Os09g23640.1 (Figure 1a; http://www.orygenesdb.cirad. fr and http://www.rice.plantbiology.msu.edu). BLASTP searches identified a close homologue of STR1 in the rice genome that shared homology with M. truncatula STR2 (Zhang et al., 2010). A whole-genome transcriptomics study using an updated version of the Affymetrix gene chip also revealedSTR2tobeinducedbyAMcolonization (C.Gutjahr, R. J. H. Sawers, H. Angliker, T. Roloff, E. Oakeley, U. Pasz- kowski, unpublished data). The STR2 transcript contained two exons, confirming the computational prediction model, jigsaw_7.642, available on the rice genome browser (http:// www.rapdb.dna.affrc.go.jp). Database analyses indicated that STR1 was accompanied by a natural antisense transcript of 1842 nt in length (cis- NAT; RAP ID, Os09t0401200-02), termed antiSTR1 hereafter. According to a full-length cDNA clone (AK106846), antiSTR1 contains two exons and is complementary with STR1 in a tail-to-tail orientation: the second exon of antiSTR1 overlaps with the third intron of STR1 by 80 nt, and with its fourth exon by 819 nt (Figure 1a). The first exon of antiSTR1 matches 131 nt of another gene adjacent to STR1 (RAP ID, Os09t0401200-01; data not shown) in sense orientation. AntiSTR1 is predicted (http://www.rapdb.dna.affrc.go.jp) to encode a protein of 94 amino acids. However, no corre- sponding protein sequence has been reported from any organism, as determined by BLAST searches. Therefore, antiSTR1 is not likely to encode for a protein but instead represents a bona fide cis-NAT. Computational searches for similar cis-NAT sequences from other plant species did not yield any match. The presence of antiSTR1 was confirmed by real-time RT-PCR, with primers specifically targeting antiSTR1 and with cDNA synthesized by an antiSTR1- specific primer (Tables S1 and S3). Natural antisense transcripts have been proposed to regulate complementary mRNA in many different ways, such as transcriptional interference, RNA masking, chro- matin remodelling, RNA editing and RNA interference (Lapidot and Pilpel, 2006). For RNA interference, small interfering (si) RNAs of mostly 21–25 nt in length are produced from the antisense transcript, and lead to Argo- naut-dependent degradation of mRNAs (Okamura and Lai, 2008). We considered the possibility that siRNAs might be produced from antiSTR1 and could also target other genes in trans. Applying a cut-off of 21-nt sequence complemen- tarity revealed a matching sequence of 121 nt in length contained within the first exon of the STR2 open reading frame, indicating the possibility that STR2 could be targeted by antiSTR1 in trans. However, as antiSTR1 originates from the same locus as STR1 we kept the name antiSTR1. Small complementary stretches of 23–34 nucleotides were also found in three other genes located on chromosomes 4 and 7, and encoding an inorganic pyrophosphatase, a glycosyl- transferase and a hypothetical protein (Table S2). However, these genes are not regulated by AM colonization (Gu ¨imil et al., 2005; C. Gutjahr, R. J. H. Sawers, H. Angliker, T. Roloff, E. Oakeley, U. Paszkowski unpublished data). Next, to confirm the AM-responsive induction of STR1, and to assess AM induction of STR2 and antiSTR1, real-time RT-PCR analysis was performed on cDNA from non-colo- nized and Glomus intraradices-colonized roots. High levels of STR1 and STR2 mRNA were detected in mycorrhizal roots, thereby confirming previous microarray data, whereas expression was at the background level in non- colonized roots (Gu ¨imil et al., 2005; C. Gutjahr, R. J. H. Sawers, H. Angliker, T. Roloff, E. Oakekly, U. Paszkowski, unpublished data). antiSTR1 mRNA also accumulated in roots, but the level did not change upon AM colonization (Figure 1b). Examination of the transcript levels of STR1, STR2 and antiSTR1 in panicle, leaf, stem and embryo revealed only background levels of expression in these organs for all three genes. As STR1 and STR2 were induced in mycorrhizal roots we next examined their expression in arbusculated cells collected by laser microdissection (Figure 1c). Transcript levels were compared between arbusculated and non- arbusculated cortex cells from mycorrhizal roots (hereafter called systemic cortex cells) and cortex cells from mock- inoculated roots. To ensure the specificity of the samples, we first recorded the expression of the arbuscule marker PT11. PT11 was specifically expressed in arbusculated cells, but not in systemic or mock control cells, as reported earlier (Gutjahr et al., 2008). Similarly, STR1 and STR2 were specifically expressed in arbusculated cortex cells. Arbus- cule-specific expression was also observed for antiSTR1. This cell-specific expression pattern was unexpected as the expression level of antiSTR1 was similar between mock and Glomus intraradices-inoculated root systems (Figure 1b). antiSTR1 transcript accumulation might therefore differ spatially between colonized and non-colonized roots, but the total level of antiSTR1 transcripts in the root system might not differ between the two treatments. In summary,
all three transcripts accumulate in arbusculated cells, suggesting they play a role in arbuscule development, maintenance or function.
Evolution
Please input evolution information here.
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Labs working on this gene
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References
Please input cited references here. Gu ¨imil, S., Chang, H., Zhu, T. et al. (2005) Comparative transcriptomics of rice reveals an ancient pattern of response to microbial colonization. Proc. Natl Acad. Sci. USA, 102, 8066–8070. Zhang, Q., Blaylock, L. and Harrison, M. (2010) Two half-ABC transporters are essential for arbuscule development in arbuscular mycorrhizal symbiosis. Plant Cell, 22, 1483–1497. Lapidot, M. and Pilpel, Y. (2006) Genome-wide natural antisense transcrip- tion: coupling its regulation to its different regulatory mechanisms. EMBO Rep. 7, 1216–1222. Gutjahr, C., Banba, M., Croset, V., An, K., Miyao, A., An, G., Hirochika, H., Imaizumi-Anraku, H. and Paszkowski, U. (2008) Arbuscular mycorrhiza- specific signaling in rice transcends the common symbiosis signaling pathway. Plant Cell, 20, 2989–3005.
Structured Information
| Gene Name |
Os09g0401100 |
|---|---|
| Description |
ABC transporter related domain containing protein |
| Version |
NM_001069654.2 GI:297609427 GeneID:4346998 |
| Length |
3671 bp |
| Definition |
Oryza sativa Japonica Group Os09g0401100, 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 9:14700465..14704135 |
| Sequence Coding Region |
14700465..14700986,14701363..14701542,14701785..14702039,14702597..14704135 |
| Expression | |
| Genome Context |
<gbrowseImage1> name=NC_008402:14700465..14704135 source=RiceChromosome09 preset=GeneLocation </gbrowseImage1> |
| Gene Structure |
<gbrowseImage2> name=NC_008402:14700465..14704135 source=RiceChromosome09 preset=GeneLocation </gbrowseImage2> |
| Coding Sequence |
<cdnaseq>atgcagcagcagcagcgcagccgtgagatggccagggccggcgagccagccgccgcccaccgccaccgcgccgagagggccgccgccggcgagccggccacccgaactcaccggacagagagagccggcgagcctgccgcggccaccaccaccaccaccacccgccggccgccgccgacgacgacggagaggaaggagagcctggagagcctcctggacgcgacggacgccgcccgtggcggccgtcgcggcggcggcggcgtgaaggcggcggtggccagcaggcaagggctggagttcaagaacctgtcgtacagcgtggtgaagaagcagaagaaggagggggtgaaggtgaagaaggaggtgtacctgctcaacgacatctccggcgaggccccccgcgggcaggtgacggcgatcctcggccccagcggcgccggcaagtccaccttcctcgacgccctcgccggccggatcgccaagggcagcctcgagggctccgtccgcatcgacggccgagctgtcacgacgagctacatgaagcagatttcatcctacgtgatgcaagatgatcagttgttcccgatgctgacggtgctggagacgctcactttcgcagctgaagtgaggctgcccccatcactctccagagctgagaagctcaagcgagtctgggagctcatcgatcagcttggtttgcagacaacggctcacacgtacattggggatgaagggacccgaggagtctccggtggggaacgccggagggtgtcgatcggcatagacatcatccacaaaccttccctcctcttcctcgacgagccgacctccggtctcgactccaccagcgcgcacagcgtggtggagaaggtgaaggacatcgccagaggagggagcattgtgctcatgacgattcaccagccatctttcaggatccagatgcttcttgacaggcttgagtacgatgaatcaacatcggggcttgagcctctggttgcctaccagagggacggcacgaaacccgacggcgccgcgaaaactccggtgccaagaacaccgagaacgccgcaccagaagtcggtgcagttcaggcagatccaactcaagagcaaccagttctccctcaacagcggcgccgccaatggcaacaccttctccaacttcgagtcgtcgtacaatgtcgatggcggcggcgatgacgacgatgaggattttgataattcgctcgagaggaagttgcagacgccgatgcacgccggcggtccagcttcggggtaccagccgaggctggcgtcacagttctacaaggatttctcggtctgggtctaccatggcgtcaccgggagcacgccgcaccggcggccgacatggacaccggcgagaacgccggtgtcgagcttccagcgtggccgcgccgtgacaatgacgccaacgccgcagaacaacccgcagcggcggcctcctccgcctccgtctccacatgtgccggtgttcaagccggaggagccgacgtaccacgagtacgagcttgacctggagccgccgctggacgcgccggaagaggactacaacggcggccatcgtcccaagttcgccaacccgtggccgcgggaggtggcggtgctgtcgtggcgcacggtgctcaacgtggtgcgcacgccggagctgttcctctctagggaggtggtgctcgcggcgatggcggtcatcctctccaccatgttccgccgcctcggcgccggcgacgtcccgacggtgaaccggctgctcaacttctacatcttcgccgtctgcctcgtcttcttctcctccaacgacgccgtgccaaccttcatccaggagcggttcatcttcatccgggagcggtcgcacaacgcctacagggcgtcgtcctacgtcgtcgcctccctcgtcgtctacctccccttcttcgccgtgcaagggctcaccttcgccgtgatcaccaagctgatgctccggatggagagcagcctgctccacttctgggtcatcctcttcgcctccctcatcaccaccaacgcctacgtcatgctcgtctcggcgctcgtccccagctacatcaccggctacgccgtcgtcatcgcgacgacggcgctcttcttcctcacctgcggcttcttcctcaagcggacgctcatcccggtggggtggaggtggctgcactacgcctccgcgatcaagtacccgttcgaggcgctgctggtgagcgagttcaaggggggccggtgctacgccggcgaccgcgccgacctctcgccggggccgctcggggggttcaagcccagcagcctccggcgggagctcaacgccagcgacgcggcgtgcccgttgatggggcaggacgtgctctccacactggacatcaccattgacagcatctgggttgatgttgccatcctgcttgcatggggtgtcctctatcgcctcctcttctatgttgtcttgaggttctactccaagaatgagaggaagtga</cdnaseq> |
| Protein Sequence |
<aaseq>MQQQQRSREMARAGEPAAAHRHRAERAAAGEPATRTHRTERAGE PAAATTTTTTRRPPPTTTERKESLESLLDATDAARGGRRGGGGVKAAVASRQGLEFKN LSYSVVKKQKKEGVKVKKEVYLLNDISGEAPRGQVTAILGPSGAGKSTFLDALAGRIA KGSLEGSVRIDGRAVTTSYMKQISSYVMQDDQLFPMLTVLETLTFAAEVRLPPSLSRA EKLKRVWELIDQLGLQTTAHTYIGDEGTRGVSGGERRRVSIGIDIIHKPSLLFLDEPT SGLDSTSAHSVVEKVKDIARGGSIVLMTIHQPSFRIQMLLDRLEYDESTSGLEPLVAY QRDGTKPDGAAKTPVPRTPRTPHQKSVQFRQIQLKSNQFSLNSGAANGNTFSNFESSY NVDGGGDDDDEDFDNSLERKLQTPMHAGGPASGYQPRLASQFYKDFSVWVYHGVTGST PHRRPTWTPARTPVSSFQRGRAVTMTPTPQNNPQRRPPPPPSPHVPVFKPEEPTYHEY ELDLEPPLDAPEEDYNGGHRPKFANPWPREVAVLSWRTVLNVVRTPELFLSREVVLAA MAVILSTMFRRLGAGDVPTVNRLLNFYIFAVCLVFFSSNDAVPTFIQERFIFIRERSH NAYRASSYVVASLVVYLPFFAVQGLTFAVITKLMLRMESSLLHFWVILFASLITTNAY VMLVSALVPSYITGYAVVIATTALFFLTCGFFLKRTLIPVGWRWLHYASAIKYPFEAL LVSEFKGGRCYAGDRADLSPGPLGGFKPSSLRRELNASDAACPLMGQDVLSTLDITID SIWVDVAILLAWGVLYRLLFYVVLRFYSKNERK</aaseq> |
| Gene Sequence |
<dnaseqindica>1..522#899..1078#1321..1575#2133..3671#atgcagcagcagcagcgcagccgtgagatggccagggccggcgagccagccgccgcccaccgccaccgcgccgagagggccgccgccggcgagccggccacccgaactcaccggacagagagagccggcgagcctgccgcggccaccaccaccaccaccacccgccggccgccgccgacgacgacggagaggaaggagagcctggagagcctcctggacgcgacggacgccgcccgtggcggccgtcgcggcggcggcggcgtgaaggcggcggtggccagcaggcaagggctggagttcaagaacctgtcgtacagcgtggtgaagaagcagaagaaggagggggtgaaggtgaagaaggaggtgtacctgctcaacgacatctccggcgaggccccccgcgggcaggtgacggcgatcctcggccccagcggcgccggcaagtccaccttcctcgacgccctcgccggccggatcgccaagggcagcctcgagggctccgtccgcatcgacggccgagctgtaagatgaatcgatgatcatcaaacacccataaaattaacctacttcctctgtttccgtttcatgttacaagatgttttaactttggttaaaaccaaactactttaaatttaactaagtttaaagaaaaaataataacattttcaaccttagacaaatttattatgaaaatatatttaattattgatttaatgaaactaattcggtattataaatattactatatttgtttataaacttagttaaaaattagtttaactttgacctaagtcaaatcgtcttgtaacttgaaacgaagagagtatacagttctctcgttcttctgcataatcaagtacacaattaagatgatctcttgttcttgttgttaatttaggtcacgacgagctacatgaagcagatttcatcctacgtgatgcaagatgatcagttgttcccgatgctgacggtgctggagacgctcactttcgcagctgaagtgaggctgcccccatcactctccagagctgagaagctcaagcgagtctgggagctcatcgatcagcttggtttgcaggtaagtcaatcaaattaatccgatagagcagttttccgatacttgagaatgtaccaggtccaatttaataccgagagataccaaattttacattaaaaaaatacggtatccttagaaccgtaaaaatctctctaatctgaccattaaacaaataatatcgacagaattgtagctggttatactctacaagctaggttagcagcttgctcatttttactgacaatctgtgtgccaaattgcagacaacggctcacacgtacattggggatgaagggacccgaggagtctccggtggggaacgccggagggtgtcgatcggcatagacatcatccacaaaccttccctcctcttcctcgacgagccgacctccggtctcgactccaccagcgcgcacagcgtggtggagaaggtgaaggacatcgccagaggagggagcattgtgctcatgacgattcaccagccatctttcaggatccagatgcttcttgacaggcttgtcatcctcgcaaggtgacaaaactatttctccctccgtttcaggttataagatgttttgactttggtcaaagtcaaactgcttcaagtttgattaagtttatagaaaaaatagtaatattttttactcatgataaatttattattaaaatatatttaattattaatttaataaaattattttggtaatataaatattactatattcgtctataaacttggtcaaacttaaaacagtttgactttcaccaaagttaaaaccttatagactgaaacgaaacgagttatagatagtattccaaattgttagtgtctaattttaaaagtttgaccaaatcttgtctgaatgaccggaggaagtatctgcctaggtctagaattgtcttttgcttccaaaattccttttttgttcaatttaaaaatactgtcattgattcatccaggggaaggttgatctacctaggaagcccaagcacactgcaaacacacctggctggatttggcaggccggtgccggatggtgagaacagcattgagtatctcctggatgtcatcaaggagtacgatgaatcaacatcggggcttgagcctctggttgcctaccagagggacggcacgaaacccgacggcgccgcgaaaactccggtgccaagaacaccgagaacgccgcaccagaagtcggtgcagttcaggcagatccaactcaagagcaaccagttctccctcaacagcggcgccgccaatggcaacaccttctccaacttcgagtcgtcgtacaatgtcgatggcggcggcgatgacgacgatgaggattttgataattcgctcgagaggaagttgcagacgccgatgcacgccggcggtccagcttcggggtaccagccgaggctggcgtcacagttctacaaggatttctcggtctgggtctaccatggcgtcaccgggagcacgccgcaccggcggccgacatggacaccggcgagaacgccggtgtcgagcttccagcgtggccgcgccgtgacaatgacgccaacgccgcagaacaacccgcagcggcggcctcctccgcctccgtctccacatgtgccggtgttcaagccggaggagccgacgtaccacgagtacgagcttgacctggagccgccgctggacgcgccggaagaggactacaacggcggccatcgtcccaagttcgccaacccgtggccgcgggaggtggcggtgctgtcgtggcgcacggtgctcaacgtggtgcgcacgccggagctgttcctctctagggaggtggtgctcgcggcgatggcggtcatcctctccaccatgttccgccgcctcggcgccggcgacgtcccgacggtgaaccggctgctcaacttctacatcttcgccgtctgcctcgtcttcttctcctccaacgacgccgtgccaaccttcatccaggagcggttcatcttcatccgggagcggtcgcacaacgcctacagggcgtcgtcctacgtcgtcgcctccctcgtcgtctacctccccttcttcgccgtgcaagggctcaccttcgccgtgatcaccaagctgatgctccggatggagagcagcctgctccacttctgggtcatcctcttcgcctccctcatcaccaccaacgcctacgtcatgctcgtctcggcgctcgtccccagctacatcaccggctacgccgtcgtcatcgcgacgacggcgctcttcttcctcacctgcggcttcttcctcaagcggacgctcatcccggtggggtggaggtggctgcactacgcctccgcgatcaagtacccgttcgaggcgctgctggtgagcgagttcaaggggggccggtgctacgccggcgaccgcgccgacctctcgccggggccgctcggggggttcaagcccagcagcctccggcgggagctcaacgccagcgacgcggcgtgcccgttgatggggcaggacgtgctctccacactggacatcaccattgacagcatctgggttgatgttgccatcctgcttgcatggggtgtcctctatcgcctcctcttctatgttgtcttgaggttctactccaagaatgagaggaagtga</dnaseqindica> |
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