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Information on pGKB5 vector for vacuum-infiltration of Arabidopsis

construction of pGKB5 - sequence

 

pGKB5 was originally described in :

Bouchez D, Camilleri C, Caboche M (1993). A binary vector based on Basta resistance for in planta transformation of Arabidopsis thaliana. C R Acad Sci Paris, Life sciences 316 : 1188-1193.

Plasmid pGKB5 was designed as a transformation vector for T-DNA tagging and is NOT a cloning vector. The T-DNA does not contain any unique cloning site, due to the large replication origin from pRiA4 which contains sites for most usual enzymes (NotI, XbaI, BamHI, XhoI, EcoRI, HindIII, PstI, SmaI, EcoRV, NruI, AccI, SalI, MluI) [5].

With the infiltration technique, stability of the replicon in the agrobacteria does not seem so crucial and any binary vector system should be usable. We recommend however the use of the C58-derived helper strain C58C1(pMP90) [6] which in our hands gave the best results for all the in planta transformation techniques (seed infection, plant decapitation, vacuum infiltration).

Construction of pGKB5

A new plant transformation vector was designed, that could be used for T-DNA tagging in Arabidopsis. The T-DNA region of this plasmid is flanked by fragments containing the right and left borders of the TR-DNA of pRiA4 [1]. The GUS-nos3' reporter cassette from pBI101.1 [2] is inserted 40 bp away from the right border . As the GUS gene possesses its own ATG initiation codon, it is able to produce active transcriptional or translational gene fusions upon its insertion in the genome. The T-DNA also contains two plant selection markers derived from pGSFR280 [3] that confer resistance of plant cells to kanamycin and to the herbicide Basta (phosphinothricin).

This binary plasmid derives from a pBGS plasmid [4], harbours a bacterial kanamycin resistance gene, and is able to replicate both in Escherichia coli and in Agrobacterium. The origin of replication of pRiA4, cloned as a large 8 kb BamHI fragment from pLJbB11 [5], confers a very high stability in Agrobacterium under non selective conditions : Agrobacterium strains containing pGKB5 show no detectable loss of the plasmid after 25 generations in medium lacking kanamycin. The binary vector was introduced into several Agrobacterium desarmed strains by electroporation : C58C1 (pMP90) [6], C58C1 (pGV2260) [7], LBA4404 [8] to give respectively the strains MP5-1, GV5-2, LB5-1.

  1. Jouanin L., Bouchez D., Drong R. F., Tepfer D., Slightom J. L. 1989. Analysis of TR-DNA / plant junctions in the genome of a Convolvulus arvensis clone transformed by Agrobacterium rhizogenes strain A4. Plant Mol. Biol. 12: 75-85.
  2. Jefferson R. A., Kavanagh T. A., Bevan M. W. 1987. GUS fusions : beta-glucuronidase as a sensitive and versatile gene fusion marker in higher plants. EMBO J. 6: 3901-3907.
  3. De Blok M., Botterman J., Vanderwiele M., Dockx J., Thoen C., GosselZ¹ V., Rao Movva N., Thompson C., Van Montagu M., Leemans J. 1987. Engineering herbicide resistance in plants by expression of a detoxifying enzyme. EMBO J. 6: 2513-2518.
  4. Spratt B. G., Hedge P. J., te Heesen S., Edelman A., Broome-Smith J. K. 1986. Kanamycin-resistant vectors that are analogs of plasmids pUC8, pUC9, pEMBL8 and pEMBL9. Gene 41: 337-342.
  5. Jouanin L., Vilaine F., d'Enfert C., Casse-Delbart F. 1985. Localization and restriction maps of the replication origin regions of the plasmids of Agrobacterium rhizogenes strain A4. Mol. Gen. Genet. 201: 370-374.
  6. Koncz C., Schell J. 1986. The promoter of TL-DNA gene 5 controls the tissue-specific expression of chimaeric genes carried by a novel type of Agrobacterium binary vector. Mol. Gen. Genet. 204: 383-396.
  7. Deblaere R., Bytebier B., De Greve H., Deboeck F., Schell J., Van Montagu M., Leemans J. 1985. Efficient octopine Ti plasmid vectors for Agrobacterium-mediated gene transfer to plants. Nucl. Acids Res. 13: 4777-4788.
  8. Hoekema A., Hirsch P. R., Hooykas P. J. J., Schilperoort R. A. 1983. A binary plant vector strategy based on separation of vir- and T-region of the Agrobacterium tumefaciens Ti plasmid. Nature 303: 179-180.

 

Sequence of pGKB5 (reconstructed & partial)

Note about the pRiA4 replication origin :
In pGKB5, the 8 kb replication origin from pRiA4 is inserted in the NotI site of the polylinker. This is a BamHI fragment with NotI linkers on both ends (and other sites, see below).
The fragment is not fully sequenced, only the BamHI-HindIII 4.7 kb fragment from the pRiA4 origin of replication has been sequenced (Mol Gen Genet 206 : 1-8 (1987)). Genbank acc # X04833, ATPRIREP.
The following restriction sites allow orientation of the fragment :
NotI,XbaI <<BamHI 0 ; XhoI 0.4 kb ; XhoI 2.05 kb ; EcoRI 3.6 kb ; XhoI 4.5 kb ; HindIII 4.7 kb ; PstI 5.3 kb ; XhoI 5.7 kb ; XhoI 6.2 kb ; BamHI 8.1 kb>>SmaI, PstI, EcoRI, EcoRV, HindIII, NruI, NotI

Reconstructed pGKB5 Sequence
(lacking the unsequenced HindIII-BamHI 3.4 kb fragment from pRiA4 origin, replaced by a run of 100 "A" in the sequence below)

ORIA4 (no sequence) + linker 1-104
Right Border (RiA4 TR-DNA) 105-622
24 bp sequence 574-596
GUS coding sequence 638-2446
NOS terminator 2505-2766
OCS terminator 2767-3489
KanR (nptII)  3490-4479
NOS promoter  4480-4766
35S promoter 4767-5924
BastaR (bar) 5925-6513
g7 terminator 6514-6768
Left Border (RiA4 TR-DNA) 6790-7525
24 bp sequence 6963-6986
Bluescript polylinker  7526-7599
pBGS vector  7599-11978
ORIA4 (4.7 kb BamHI-HindIII sequenced) 11979-16615
ORIA4 (no sequence) 16616-16686

AAAAAAAAAAAAAAAAAAAAAAAAAAAAAAAGGATCCCCCGGGCTGCAGG
AATTCGATATCAAGCTTATCGCGATACCGTCGGCGGCCGC
TCTAGAGGAT
CCCCCCACAGACAGCTCCGTAGCCCTCGTTCTCCTTGGAGTTCTTCGGGA
AATGGATCTTTCGATTCCCGATGATGTCTCTCTTATCTGCTTTGACGACG
CCGACTGGACATCCGCTATAACGCCGCCATTGACCGTGATTTCGCAACCT
GTCAGGGATCTCGCGACGGCTGCCACAGAAGACCTGATCGCCCGCTTAAA
GGGCGAGACTTCAGCCCCACCCAAGGAAACTCTTCTCCCGGCGGTTCTCA
TAGAGCGCGGTTCCGTAAGCGGTTCTTCGCAAGGTCGGGGTTGCATACCG
AACTCGCGAAACGTCGGCGACTGAGCTCCCGAGGCGCGTTGACAAGATGC
CACGAAGGGAATGGAAGACAGCCGATATTGCAATTGTCTTCGTGGACTGC
TTTCGGGACGTAAGGCGCAAGCCATCATCACCGCCGTCCTAAACAAACAT
ACCTCCACACAAATTTATCTACCTGACCACAAGATATATCCTGTCACACG
ATTTATTAAACGCTGCACTTGGGGTGGTCAGTCCCTTATGTTACGTCCTG
TAGAAACCCCAACCCGTGAAATCAAAAAACTCGACGGCCTGTGGGCATTC
AGTCTGGATCGCGAAAACTGTGGAATTGATCAGCGTTGGTGGGAAAGCGC
GTTACAAGAAAGCCGGGCAATTGCTGTGCCAGGCAGTTTTAACGATCAGT
TCGCCGATGCAGATATTCGTAATTATGCGGGCAACGTCTGGTATCAGCGC
GAAGTCTTTATACCGAAAGGTTGGGCAGGCCAGCGTATCGTGCTGCGTTT
CGATGCGGTCACTCATTACGGCAAAGTGTGGGTCAATAATCAGGAAGTGA
TGGAGCATCAGGGCGGCTATACGCCATTTGAAGCCGATGTCACGCCGTAT
GTTATTGCCGGGAAAAGTGTACGTATCACCGTTTGTGTGAACAACGAACT
GAACTGGCAGACTATCCCGCCGGGAATGGTGATTACCGACGAAAACGGCA
AGAAAAAGCAGTCTTACTTCCATGATTTCTTTAACTATGCCGGAATCCAT
CGCAGCGTAATGCTCTACACCACGCCGAACACCTGGGTGGACGATATCAC
CGTGGTGACGCATGTCGCGCAAGACTGTAACCACGCGTCTGTTGACTGGC
AGGTGGTGGCCAATGGTGATGTCAGCGTTGAACTGCGTGATGCGGATCAA
CAGGTGGTTGCAACTGGACAAGGCACTAGCGGGACTTTGCAAGTGGTGAA
TCCGCACCTCTGGCAACCGGGTGAAGGTTATCTCTATGAACTGTGCGTCA
CAGCCAAAAGCCAGACAGAGTGTGATATCTACCCGCTTCGCGTCGGCATC
CGGTCAGTGGCAGTGAAGGGCCAACAGTTCCTGATTAACCACAAACCGTT
CTACTTTACTGGCTTTGGTCGTCATGAAGATGCGGACTTACGTGGCAAAG
GATTCGATAACGTGCTGATGGTGCACGACCACGCATTAATGGACTGGATT
GGGGCCAACTCCTACCGTACCTCGCATTACCCTTACGCTGAAGAGATGCT
CGACTGGGCAGATGAACATGGCATCGTGGTGATTGATGAAACTGCTGCTG
TCGGCTTTAACCTCTCTTTAGGCATTGGTTTCGAAGCGGGCAACAAGCCG
AAAGAACTGTACAGCGAAGAGGCAGTCAACGGGGAAACTCAGCAAGCGCA
CTTACAGGCGATTAAAGAGCTGATAGCGCGTGACAAAAACCACCCAAGCG
TGGTGATGTGGAGTATTGCCAACGAACCGGATACCCGTCCGCAAGTGCAC
GGGAATATTTCGCCACTGGCGGAAGCAACGCGTAAACTCGACCCGACGCG
TCCGATCACCTGCGTCAATGTAATGTTCTGCGACGCTCACACCGATACCA
TCAGCGATCTCTTTGATGTGCTGTGCCTGAACCGTTATTACGGATGGTAT
GTCCAAAGCGGCGATTTGGAAACGGCAGAGAAGGTACTGGAAAAAGAACT
TCTGGCCTGGCAGGAGAAACTGCATCAGCCGATTATCATCACCGAATACG
GCGTGGATACGTTAGCCGGGCTGCACTCAATGTACACCGACATGTGGAGT
GAAGAGTATCAGTGTGCATGGCTGGATATGTATCACCGCGTCTTTGATCG
CGTCAGCGCCGTCGTCGGTGAACAGGTATGGAATTTCGCCGATTTTGCGA
CCTCGCAAGGCATATTGCGCGTTGGCGGTAACAAGAAAGGGATCTTCACT
CGCGACCGCAAACCGAAGTCGGCGGCTTTTCTGCTGCAAAAACGCTGGAC
TGGCATGAACTTCGGTGAAAAACCGCAGCAGGGAGGCAAACAATGAATCA
ACAACTCTCCTGGCGCACCATCGTCGGCTACAGCCTCGGTGGGGAATTGA
GCTCGATCGTTCAAACATTTGGCAATAAAGTTTCTTAAGATTGAATCCTG
TTGCCGGTCTTGCGATGATTATCATATAATTTCTGTTGAATTACGTTAAG
CATGTAATAATTAACATGTAATGCATGACGTTATTTATGAGATGGGTTTT
TATGATTAGAGTCCCGCAATTATACATTTAATACGCGATAGAAAACAAAA
TATAGCGCGCAAACTAGGATAAATTATCGCGCGCGGTGTCATCTATGTTA
CTAGATCGAATTCGATCGAGGGGATCGAGCCCCTGCTGAGCCTCGACATG
TTGTCGCAAAATTCGCCCTGGACCCGCCCAACGATTTGTCGTCACTGTCA
AGGTTTGACCTGCACTTCATTTGGGGCCCACATACACCAAAAAAATGCTG
CATAATTCTCGGGGCAGCAAGTCGGTTACCCGGCCGCCGTGCTGGACCGG
GTTGAATGGTGCCCGTAACTTTCGGTAGAGCGGACGGCCAATACTCAACT
TCAAGGAATCTCACCCATGCGCGCCGGCGGGGAACCGGAGTTCCCTTCAG
TGAACGTTATTAGTTCGCCGCTCGGTGTGTCGTAGATACTAGCCCCTGGG
GCCTTTTGAAATTTGAATAAGATTTATGTAATCAGTCTTTTAGGTTTGAC
CGGTTCTGCCGCTTTTTTTAAAATTGGATTTGTAATAATAAAACGCAATT
GTTTGTTATTGTGGCGCTCTATCATAGATGTCGCTATAAACCTATTCAGC
ACAATATATTGTTTTCATTTTAATATTGTACATATAAGTAGTAGGGTACA
ATCAGTAAATTGAACGGAGAATATTATTCATAAAAATACGATAGTAACGG
GTGATATATTCATTAGAATGAACCGAAACCGGCGGTAAGGATCTGAGCTA
CACATGCTCAGGTTTTTTACAACGTGCACAACAGAATTGAAAGCAAATAT
CATGCGATCATAGGCGTCTCGCATATCTCATTAAAGCAGGGGGTGGGCGA
AGAACTCCAGCATGAGATCCCCGCGCTGGAGGATCATCCAGCCGGCGTCC
CGGAAAACGATTCCGAAGCCCAACCTTTCATAGAAGGCGGCGGTGGAATC
GAAATCTCGTGATGGCAGGTTGGGCGTCGCTTGGTCGGTCATTTCGAACC
CCAGAGTCCCGCTCAGAAGAACTCGTCAAGAAGGCGATAGAAGGCGATGC
GCTGCGAATCGGGAGCGGCGATACCGTAAAGCACGAGGAAGCGGTCAGCC
CATTCGCCGCCAAGCTCTTCAGCAATATCACGGGTAGCCAACGCTATGTC
CTGATAGCGGTCCGCCACACCCAGCCGGCCACAGTCGATGAATCCAGAAA
AGCGGCCATTTTCCACCATGATATTCGGCAAGCAGGCATCGCCATGGGTC
ACGACGAGATCCTCGCCGTCGGGCATGCGCGCCTTGAGCCTGGCGAACAG
TTCGGCTGGCGCGAGCCCCTGATGCTCTTCGTCCAGATCATCCTGATCGA
CAAGACCGGCTTCCATCCGAGTACGTGCTCGCTCGATGCGATGTTTCGCT
TGGTGGTCGAATGGGCAGGTAGCCGGATCAAGCGTATGCAGCCGCCGCAT
TGCATCAGCCATGATGGATACTTTCTCGGCAGGAGCAAGGTGAGATGACA
GGAGATCCTGCCCCGGCACTTCGCCCAATAGCAGCCAGTCCCTTCCCGCT
TCAGTGACAACGTCGAGCACAGCTGCGCAAGGAACGCCCGTCGTGGCCAG
CCACGATAGCCGCGCTGCCTCGTCCTGCAGTTCATTCAGGGCACCGGACA
GGTCGGTCTTGACAAAAAGAACCGGGCGCCCCTGCGCTGACAGCCGGAAC
ACGGCGGCATCAGAGCAGCCGATTGTCTGTTGTGCCCAGTCATAGCCGAA
TAGCCTCTCCACCCAAGCGGCCGGAGAACCTGCGTGCAATCCATCTTGTT
CAATCCACATGATCATGGGCCGGATCTTTGATTGAGAGTGAATATGAGAC
TCTAATTGGATACCGAGGGGAATTTATGGAACGTCAGTGGAGCATTTTTG
ACAAGAAATATTTGCTAGCTGATAGTGACCTTAGGCGACTTTTGAACGCG
CAATAATGGTTTCTGACGTATGTGCTTAGCTCATTAAACTCCAGAAACCC
GCGGCTGAGTGGCTCCTTCAATCGTTGCGGTTCTGTCAGTTCCAAACGTA
AAACGGCTTGTCCCGCGTCATCGGCGGGGGTCATAACGTGACTCCCTTAA
TTCTCCGCTCATGATCCTGTTTCCTGTGTGAAATTGTTATCCGCTCACAA
TTCCACACATTATACGAGCCGGAAGCATAAAGTGTAAAGCCTGGGGTGCC
TAATGAGTGAGCTAACTCACATTAATTGCGTTGCGCTCACTGCCCGCTTT
CCAGTCGGGAAACCTGTCGTGCCAGCTGCATTAATGAATCGGAATTGACG
GATCTCCTTTGCCCCGGAGATCACCATGGACGACTTTCTCTATCTCTACG
ATCTAGGAAGAAAGTTCGACGGAGAAGGTGACGATACCATGTTCACCACC
GATAATGAGAAGATTAGCCTCTTCAATTTCAGAAAGAATGCTGACCCACA
GATGGTTAGAGAGGCCTACGCGGCAGGTCTCATCAAGACGATCTACCCGA
GTAATAATCTCCAGGAGATCAAATACCTTCCCAAGAAGGTTAAAGATGCA
GTCAAAAGATTCAGGACTAACTGCATCAAGAACACAGAGAAAGATATATT
TCTCAAGATCAGAAGTACTATTCCAGTATGGACGATTCAAGGCTTGCTTC
ATAAACCAAGGCAAGTAATAGAGATTGGAGTCTCTAAGAAAGTAGTTCCT
ACTGAATCAAAGGCCATGGAGTCAAAAATTCAGATCGAGGATCTAACAGA
ACTCGCCGTGAAGACTGGCGAACAGTTCATACAGAGTCTTTTACGACTCA
ATGACAAGAAGAAAATCTTCGTCAACATGGTGGAGCACGACACTCTCGTC
TACTCCAAGAATATCAAAGATACAGTCTCAGAAGACCAAAGGGCTATTGA
GACTTTTCAACAAAGGGTAATATCGGGAAACCTCCTCGGATTCCATTGCC
CAGCTATCTGTCACTTCATCAAAAGGACAGTAGAAAAGGAAGGTGGCACC
TACAAATGCCATCATTGCGATAAAGGAAAGGCTATCGTTCAAGATGCCTC
TGCCGACAGTGGTCCCAAAGATGGACCCCCACCCACGAGGAGCATCGTGG
AAAAAGAAGACGTTCCAACCACGTCTTCAAAGCAAGTGGATTGATGTGAT
ATCTCCACTGACGTAAGGGATGACGCACAATCCCACTATCCTTCGCAAGA
CCCTTCCTCTATATAAGGAAGTTCATTTCATTTGGAGAGGACACGCTGAA
ATCACCAGTCTCTCTCTACAAATCGGATCCATGAGCCCAGAACGACGCCC
GGCCGACATCCGCCGTGCCACCGAGGCGGACATGCCGGCGGTCTGCACCA
TCGTCAACCACTACATCGAGACAAGCACGGTCAACTTCCGTACCGAGCCG
CAGGAACCGCAGGAGTGGACGGACGACCTCGTCCGTCTGCGGGAGCGCTA
TCCCTGGCTCGTCGCCGAGGTGGACGGCGAGGTCGCCGGCATCGCCTACG
CGGGCCCCTGGAAGGCACGCAACGCCTACGACTGGACGGCCGAGTCGACC
GTGTACGTCTCCCCCCGCCACCAGCGGACGGGACTGGGCTCCACGCTCTA
CACCCACCTGCTGAAGTCCCTGGAGGCACAGGGCTTCAAGAGCGTGGTCG
CTGTCATCGGGCTGCCCAACGACCCGAGCGTGCGCATGCACGAGGCGCTC
GGATATGCCCCCCGCGGCATGCTGCGGGCGGCCGGCTTCAAGCACGGGAA
CTGGCATGACGTGGGTTTCTGGCAGCTGGACTTCAGCCTGCCGGTACCGC
CCCGTCCGGTCCTGCCCGTCACCGAGATCTGATCTCACGCGTCTAGGATC
CGATGGATCCCCCGATGAGCTAAGCTAGCTATATCATCAATTTATGTATT
ACACATAATATCGCACTCAGTCTTTCATCTACGGCAATGTACCAGCTGAT
ATAATCAGTTATTGAAATATTTCTGAATTTAAACTTGCATCAATAAATTT
ATGTTTTTGCTTGGACTATAATACCTGACTTGTTATTTTATCAATAAATA
TTTAAACTATATTTCTTTCAAGATGGGAATTAACATCTACAAATTGCCTT
TTCTTATCGACCATGTACATCAAGCTTATCGATACCGTCGGCTATTGGTA
ATAGGACACTGGGATTCGTCTTGGACAACTTTCCTTCTCATCTAAGCGTA
GACAACCCTCAACTGGAAACGGGCCGGACTCCAGGGCGTGTGCCAGGTGC
CCACGGAATAGTTTTGGCCAGACCCTTGAAAATCCGATTCAGTACAATCG
ATTGCCCTCATTTTTACGTTGGCATATATCCTGCCAAACAGCCAACAACG
CGCGTGCGGTGAATAGGAAAGCGTTTGAGTTGCTTGCTCATATCGTGACG
GTTGACAGCACAGGTTGACCGCTTGATGATTCGTACGAGCCGCCAAACAT
TGGCTGTCGTAATGATATACCATGTCAGAACAGCAATCCGATGGGGCGGA
AAGCATTATCTTAATGCACACGGAAATGGCGCGTCGGTGGGTGGAATACA
CCGACATAGAGGCCGTAAGTTCTGCATGGTCATCGTCGGAAAGGTGGCAG
CAGGCGCACGGCTGTGGCCTCTTGCTCTTTCAGCGTGAAATGCGTGTTGA
AAGAATAATCGAAGAGAGCGTCCGCTCGACACCTTCAATTATGCCGATTT
GATCGATGAACTGATCGAGCTCTGAAATCGAAGGGGCTTCGATAATCGCA
ATCAAATCAAAAGTGCCACTCACAGAATGAAGAGCGATAACGGCCGTGAC
CTTCCCAAGGGAGGCCGTCACCTGTGAAAGCGCCTTCGTAATGGTGATCA
GAATATGGGCTCGAACCAAGCTCGAGACCTCGAGGGGGGGCCCGGTACCC
AATTCGCCCTATAGTGAGTCGTATTACAATTCACTGGCCGTCGTTTTACA
ACGTCGTGACTGGGAAAACCCTGGCGTTACCCAACTTAATCGCCTTGCAG
CACATCCCCCTTTCGCCAGCTGGCGTAATAGCGAAGAGGCCCGCACCGAT
CGCCCTTCCCAACAGTTGCGCAGCCTGAATGGCGAATGCCCCTGTAATCC
GGGCAGCGCAACGGAACATTCATCAGTGTAAAAATGGAATCAATAAAGCC
CTGCGCAGCGCGCAGGGTCAGCCTGAATACGCGTTTAATGACCAGCACAG
TCGTGATGGCAAGGTCAGAATAGCGCTGAGGTCTGCCTCGTGAAGAAGGT
GTTGCTGACTCATACCAGGCCTGAATCGCCCCATCATCCAGCCAGAAAGT
GAGGGAGCCACGGTTGATGAGAGCTTTGTTGTAGGTGGACCAGTTGGTGA
TTTTGAACTTTTGCTTTGCCACGGAACGGTCTGCGTTGTCGGGAAGATGC
GTGATCTGATCCTTCAACTCAGCAAAAGTTCGATTTATTCAACAAAGCCG
CCGTCCCGTCAAGTCAGCGTAATGCTCTGCCAGTGTTACAACCAATTAAC
CAATTCTGATTAGAAAAACTCATCGAGCATCAAATGAAACTGCAATTTAT
TCATATCAGGATTATCAATACCATATTTTTGAAAAAGCCGTTTCTGTAAT
GAAGGAGAAAACTCACCGAGGCAGTTCCATAGGATGGCAAGATCCTGGTA
TCGGTCTGCGATTCCGACTCGTCCAACATCAATACAACCTATTAATTTCC
CCTCGTCAAAAATAAGGTTATCAAGTGAGAAATCACCATGAGTGACGACT
GAATCCGGTGAGAATGGCAAAAGCTTATGCATTTCTTTCCAGACTTGTTC
AACAGGCCAGCCATTACGCTCGTCATCAAAATCACTCGCATCAACCAAAC
CGTTATTCATTCGTGATTGCGCCTGAGCGAGACGAAATACGCGATCGCTG
TTAAAAGGACAATTACAAACAGGAATCGAATGCAACCGGCGCAGGAACAC
TGCCAGCGCATCAACAATATTTTCACCTGAATCAGGATATTCTTCTAATA
CCTGGAATGCTGTTTTCCCGGGGATCGCAGTGGTGAGTAACCATGCATCA
TCAGGAGTACGGATAAAATGCTTGATGGTCGGAAGAGGCATAAATTCCGT
CAGCCAGTTTAGTCTGACCATCTCATCTGTAACATCATTGGCAACGCTAC
CTTTGCCATGTTTCAGAAACAACTCTGGCGCATCGGGCTTCCCATACAAT
CGATAGATTGTCGCACCTGATTGCCCGACATTATCGCGAGCCCATTTATA
CCCATATAAATCAGCATCCATGTTGGAATTTAATCGCGGCCTCGAGCAAG
ACGTTTCCCGTTGAATATGGCTCATAACACCCCTTGTATTACTGTTTATG
TAAGCAGACAGTTTTATTGTTCATGATGATATATTTTTATCTTGTGCAAT
GTAACATCAGAGATTTTGAGACACAACGTGGCTTTGTTGAATAAATCGAA
CTTTTGCTGAGTTGAAGGATCAGATCACGCATCTTCCCGACAACGCAGAC
CGTTCCGTGGCAAAGCAAAAGTTCAAAATCACCAACTGGTCCACCTACAA
CAAAGCTCTCATCAACCGTGGCTCCCTCACTTTCTGGCTGGATGATGGGG
CGATTCAGGCCTGGTATGAGTCAGCAACACCTTCTTCACGAGGCAGACCT
CAGCGCTATTCTGACCTTGCCATCACGACTGTGCTGGTCATTAAACGCGT
ATTCAGGCTGACCCTGCGCGCTGCGCAGGGCTTTATTGATTCCATTTTTA
CACTGATGAATGTTCCGTTGCGCTGCCCGGATTACAGGGGGCAATGGCAA
CAACGTTGCGCAAACTATTAACTGGCGAACTACTTACTCTAGCTTCCCGG
CAACAATTAATAGACTGGATGGAGGCGGATAAAGTTGCAGGACCACTTCT
GCGCTCGGCCCTTCCGGCTGGCTGGTTTATTGCTGATAAATCTGGAGCCG
GTGAGCGTGGGTCTCGCGGTATCATTGCAGCACTGGGGCCAGATGGTAAG
CCCTCCCGTATCGTAGTTATCTACACGACGGGGAGTCAGGCAACTATGGA
TGAACGAAATAGACAGATCGCTGAGATAGGTGCCTCACTGATTAAGCATT
GGTAACTGTCAGACCAAGTTTACTCATATATACTTTAGATTGATTTATTG
TTCTGGATATTACCAGCAAGGCCGATAGTTTGAGTTCTTCTACTCAGGCA
AGTGATGTTATTACTAATCAAAGAAGTATTGCGACAACGGTTAATTTGCG
TGATGGACAGACTCTTTTGCTCGGTGGCCTCACTGATTATAAAAACACTT
CTCAGGATTCTGGCGTACCGTTCCTGTCTAAAATCCCTTTAATCGGCCTC
CTGTTTAGCTCCCGCTCTGATTCTAACGAGGAAAGCACGTTATACGTGCT
CGTCAAAGCAACCATAGTACGCGCCCTGTAGCGGCGCATTAAGCGCGGCG
GGTGTGGTGGTTACGCGCAGCGTGACCGCTACACTTGCCAGCGCCCTAGC
GCCCGCTCCTTTCGCTTTCTTCCCTTCCTTTCTCGCCACGTTCGCCGGCT
TTCCCCGTCAAGCTCTAAATCGGGGGCTCCCTTTAGGGTTCCGATTTAGT
GCTTTACGGCACCTCGACCCCAAAAAACTTGATTAGGGTGATGGTTCACG
TAGTGGGCCATCGCCCTGATAGACGGTTTTTCGCCCTTTGACGTTGGAGT
CCACGTTCTTTAATAGTGGACTCTTGTTCCAAACTGGAACAACACTCAAC
CCTATCTCGGTCTATTCTTTTGATTTATAAGGGATTTTGCCGATTTCGGC
CTATTGGTTAAAAAATGAGCTGATTTAACAAAAATTTAACGCGAATTTTA
ACAAAATAAAAGGATCTAGGTGAAGATCCTTTTTGATAATCTCATGACCA
AAATCCCTTAACGTGAGTTTTCGTTCCACTGAGCGTCAGACCCCGTAGAA
AAGATCAAAGGATCTTCTTGAGATCCTTTTTTTCTGCGCGTAATCTGCTG
CTTGCAAACAAAAAAACCACCGCTACCAGCGGTGGTTTGTTTGCCGGATC
AAGAGCTACCAACTCTTTTTCCGAAGGTAACTGGCTTCAGCAGAGCGCAG
ATACCAAATACTGTCCTTCTAGTGTAGCCGTAGTTAGGCCACCACTTCAA
GAACTCTGTAGCACCGCCTACATACCTCGCTCTGCTAATCCTGTTACCAG
TGGCTGCTGCCAGTGGCGATAAGTCGTGTCTTACCGGGTTGGACTCAAGA
CGATAGTTACCGGATAAGGCGCAGCGGTCGGGCTGAACGGGGGGTTCGTG
CACACAGCCCAGCTTGGAGCGAACGACCTACACCGAACTGAGATACCTAC
AGCGTGAGCATTGAGAAAGCGCCACGCTTCCCGAAGGGAGAAAGGCGGAC
AGGTATCCGGTAAGCGGCAGGGTCGGAACAGGAGAGCGCACGAGGGAGCT
TCCAGGGGGAAACGCCTGGTATCTTTATAGTCCTGTCGGGTTTCGCCACC
TCTGACTTGAGCGTCGATTTTTGTGATGCTCGTCAGGGGGGCGGAGCCTA
TGGAAAAACGCCAGCAACGCGGCCTTTTTACGGTTCCTGGCCTTTTGCTG
GCCTTTTGCTCACATGTTCTTTCCTGCGTTATCCCCTGATTCTGTGGATA
ACCGTATTACCGCCTTTGAGTGAGCTGATACCGCTCGCCGCAGCCGAACG
ACCGAGCGCAGCGAGTCAGTGAGCGAGGAAGCGGAAGAGCGCCTGATGCG
GTATTTTCTCCTTACGCATCTGTGCGGTATTTCACACCGCATATGGTGCA
CTCTCAGTACAATCTGCTCTGATGCCGCATAGTTAAGCCAGTATATACAC
TCCGCTATCGCTACGTGACTGGGTCATGGCTGCGCCCCGACACCCGCCAA
CACCCGCTGACGCGCCCTGACGGGCTTGTCTGCTCCCGGCATCCGCTTAC
AGACAAGCTGTGACCGTCTCCGGGAGCTGCATGTGTCAGAGGTTTTCACC
GTCATCACCGAAACGCGCGAGGCAGCCAATACGCAAACCGCCTCTCCCCG
CGCGTTGGCCGATTCATTAATGCAGCTGGCACGACAGGTTTCCCGACTGG
AAAGCGGGCAGTGAGCGCAACGCAATTAATGTGAGTTAGCTCACTCATTA
GGCACCCCAGGCTTTACACTTTATGCTTCCGGCTCGTATGTTGTGTGGAA
TTGTGAGCGGATAACAATTTCACACAGGAAACAGCTATGACCATGATTAC
GCCAAGCTCGGAATTAACCCTCACTAAAGGGAACAAAAGCTGGAGCTCCA
CCGCGGTGGCGGCCGCTCTAGAACTAGTggatcctacaaggtagaatccg
cctgagtcgcaagggtgacttcgcctatattggacgacggcgcgcagagg
gcgacctctttttgggttacgattgtaggattatcactaaaacaatacat
gaacatattcaaatggcaatctctctaaggcattggaaataaatacaaat
aacagttgggtggagtttttcgacctgagggcgttaaccttctgttaacc
taaaagctcttgcccaaacagcagaatcggcgctaattgccagcggcgga
acttttccagtttcgcgaaaaatatcgccactggcaaggaatgggtttga
gatggcgaagtctgtcctaaaagcagcgcctgtagttgtagggttgacgg
ccttgatggagcgtcatgccgatgccctctcgagccaacttcaagcacat
catcttaaggttttcccgccgcattccgagaagggcattcgaacattcgg
gccatcggaggcgtccaagctgctcggcgttggcgagtcatatttacggc
agaccgcgtctgagatgccagagttgaatgttagcatgagcccgggtggc
aggcgaatgttctcaattgaagatatccatgtgattcggaagtatatgga
tcaggtcggccgcgggaaccggcgctacctgccacatcgtcgaggcggcg
agcagcttcaggttatctctgtgatgaatttcaaaggtgggtcgggtaag
accaccaccgccgcgcatctggcgcagtacctcgctatgcgcggatatcg
agtcttggccattgatctcgatcctcaagcgagcctttctgcactctttg
ggagccaaccggagacggacgttggcccgaacgaaacgctctacggcgct
ataaggtatgatgatgagcaggtggcaatcgaacgagtcgtccgagggac
ttacattcccgacctccacctgattcctggtaaccttgagctgatggagt
ttgaacacgatacgccacgcgcgctgatgaaccgcaaagagggcgacacg
ctcttttatggtcgcatcagccaagtaattgaagatatcgcggataacta
tgacgtcgtggtcatcgactgccctccccagcttgggtatctcacgctat
ccgcattgactgcggcgacgtccattcttgtcacggtccatccgcagatg
ctggatgtgatgtcgatgaaccagtttctggcaatgacatcgaacctttt
gcgtgaaatcgagaatgctggcgccaagttcaagtttaattggatgcgct
atctgataacccgtttcgaaccgagcgacggaccacagaaccaaatggta
ggttatctgcggtcgatttttggcgaaaatgtcctcaattttccgatgct
taaaaccaccgcggtttcggacgctggcctgacaaaccagactctattcg
aagtggagcgtggcctgttcacgcgctcgacctatgatcgagccttggag
gcgatgaacgccgtcaacgacgagatcgaaacactgatcaaaaaagcatg
gggtaggcccacatgagccggaagcacatccttggcgtctcaactgacgc
ccctgagacgtcgcccgccgacaataggacggcaaagaaccgctccatgc
cgctcctcggcgtaacaaggaaggagcgcgatccggcaacgaagctcaca
gcgaacattggtaacgcactgcgagagcaaaacgatcgtcttagccgtgc
cgaagagatcgagcggcgtctcgctgaaggtcaggcagtgatagagttgg
atgcctcgtcaatagaaccgtctttcgtgcaggatcgtatgcgaggggac
attgacgggctccttacttcgatccgggaacaaggacagcaagtcccaat
ccttgtgcgaccgcatccgagccagccgggccgatatcaggttgccttcg
gccaccgccggctacgcgccgtttcagaactcggacttccggtcagagcg
gtcgttcgcgaactgacggacgagcaagtggtcgtagcacagggtcagga
aaacaatgagcgcgaagatcttaccttcatcgaaaaggcgcgcttcgcac
atcgcctgaacaggcagttttctcgagagattgtcatcgccgcgatgtcg
atcgacaagagcaatttgtccaagatgcttctgctcgtcgacgccctccc
ctctgaactgaccgatgctattggtgccgctcctggtgttggacggccga
gttggcaacaacttgccgagctgattgagaaagtttcttcaccggccgac
gtggctaaatatgctatgtcggaggaagttcaagcgctgccatcggcaga
acgattcaaggcggtgatcgctagtctgaagcccagtcgggttgcgcgtg
gacttcccgaggtcatggccaccccagacggcaccagaattgcacaggtg
acgcagagcaaggccaaactggaaatcacgattgacaggaaggcgacgcc
cgattttgcgaccttcgtgctcgatcatgtgccagcgctgtatcaagcgt
accacgctgagaaccaacggaaacggggagagtaaaccgcaaaagaaaag
agccccctcaacgtcgccgtcgcggaagcccttctgtctctctagcgcga
acagaatcgcatttcctcgaatcctcgtcaagagtttttagcgccgtttt
ggtgagctgatttcctttgcctgctgaaaggtgaaagatgatgcagacag
gaagtgtaacgacgccattcgggcggcggccaatgacgcttgcgcttgtg
cggcgccagacggcgctggccgatatcaaacaaggcaagacagcggacaa
gtggaaggtctttagagacgcgtccgcggccatggaactacttggaatcc
agtccaacagtcttgccgtccttgatgcgctattgagctttcacccggaa
acggagttgcgtcaggaggcacagctgatcgtcttcccgtcgaatgctca
gcttgcccttcgggcgcatgggatggctggcgcgactttgcgtaggcaca
tcgccatgctcgtggagtcaggcttgatcgtccggaaggatagcgccaac
ggaaagcgttacgctcgtaaggatggcgctggtcagatcgagcgcgcgtt
tggcttcgatttgtctccgcttctcgcgcggtccgaagagctagcgatga
tggcacagcaggtgatggccgatcgagcagcattcaggatggccaaagaa
agtctgacgatttgccgacgggacgttcggaagctaattacggcagctat
ggaagagggagcggagggcgactggcaagctgtcgaggaagtctatgtgg
aacttgtgggtagaattccacgcgccccgacgcttgctgatgtagagtca
attctcgaagagatgtggatgctccaggaagagataatcaaccggttgga
aattagagacaattcagaaaataatagcaccaatgctgcccagagcgagc
agcacatacagaattcaaaacccgaatccgttaatgaacttgaacctcgc
tctgaaaaggagcagggcgctaagccgagtgaaatagaccgggcaaggag
cgagccgataaaagcgttccccctcgggatgatcctgaaagcatgcccga
ccattggcaattatgggccgagcggtgcggttgctagctggcgtgacctc
atgtcggctgcggtggtggttcggtctatgctgggggtcagcccgtcggc
ttaccaagacgcgtgtgaggcaatgggaccggagaatgcggcagcagcga
tggcgtgcattttggagcgagcgaacttcatcaattcgcccgggggctat
ctccgagatctgacacggcggagcgagctcgggaagttttcacttggccc
gatgataatggcgctcttgaaggctagcgggcaggggacgttgcggtttg
gctagaattagcgagtatggagcaggatggtctgtggtcagctgaccaca
gacctaataggttgaaaacatgagcgttttttggatgatcgacagaccat
ccgattcccggagtaccaagcgtgctctgatgggagcgataacattactc
aacaagcacgaaggccccatgccgatcgttgatcgtgaaggagagcctgc
tctacatgcggcggtattttgccggccgaggcatgtagtcgcggagcact
gcctatttactgccctaggcacaaacgttgactcttggatcgagctggca
gacaaagcaataacccacacagaggacgattaatggctgacgaagagatc
cagaatccgccggacggtactgctgctgccgaagttgagccggctgctcc
tagaggtagaagagcaaagaaagcaccagccgaaacagcccgcacgggat
cgttcaaatccgtgaagccgaaaacccgcggcctcagcaaccgagaaaaa
ctggagaagatcggtcaaatcgaagctcaggtcgctggcggcgcaacctt
gaaggacgccgttaagatcgtgggtatttccgttcagacctattatcaat
ggaagagagctgcggttcaacctgtctcacagaatccggccgtgtctgtt
tcagttgacgatgaactcggcgagttcatccaactcgaggaggaaaatcg
gcggctcagaaagctt
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