Title:
AGROBACTERIUM BACTERIUM TO BE USED IN PLANT TRANSFORMATION METHOD
Document Type and Number:
WIPO Patent Application WO/2014/157541
Kind Code:
A1
Abstract:
An Agrobacterium bacterium to be used in a plant transformation method, said Agrobacterium bacterium containing three kinds of plasmids:
(1) a plasmid comprising the following constituents
(i) genes virB, virC, virD1, virD2, virD3, virG and virJ of pTiBo542, and
(ii) a replication initiation point;
(2) a disarmed Ti plasmid or a disarmed Ri plasmid of the Agrobacterium bacterium; and
(3) a plasmid having a T-DNA region comprising a desired DNA,
wherein the plasmids (1) to (3) have replication mechanisms capable of coexisting with each other.
Inventors:
IMAYAMA TERUYUKI (JP)
HIEI YUKOH (JP)
ISHIDA YUJI (JP)
HIEI YUKOH (JP)
ISHIDA YUJI (JP)
Application Number:
PCT/JP2014/058926
Publication Date:
October 02, 2014
Filing Date:
March 27, 2014
Export Citation:
Assignee:
JAPAN TOBACCO INC (JP)
International Classes:
C12N15/09; A01H1/00; A01H5/00; C12N1/21
Domestic Patent References:
| WO2007148819A1 | 2007-12-27 | |||
| WO2007148819A1 | 2007-12-27 |
Foreign References:
| US20090075358A1 | 2009-03-19 |
Other References:
WENCK A.R. ET AL.: "High-efficiency Agrobacterium-mediated transformation of Norway spruce (Picea abies) and loblolly pine (Pinus taeda", PLANT MOL. BIOL., vol. 39, 1999, pages 407 - 416, XP002909525
HIEI Y. ET AL.: "Efficient transformation of rice (Oryza sativa L.) mediated by Agrobacterium and sequence analysis of the boundaries of the T-DNA", PLANT J., vol. 6, no. 2, August 1994 (1994-08-01), pages 271 - 282, XP002922710
AMOAH B.K. ET AL.: "Factors influencing Agrobacterium-mediated transient expression of uidA in wheat inflorescence tissue", J. EXP. BOT., vol. 52, no. 358, May 2001 (2001-05-01), pages 1135 - 1142, XP002995060
JIN, S.; KOMARI, T.; GORDON, M.P.; NESTER, E.W.: "Genes responsible for the supervirulence phenotype of Agrobacterium tumefaciens A281", J. BACTERIOL., vol. 169, 1987, pages 4417 - 4425, XP008101731, DOI: doi:10.1128/jb.169.10.4417-4425.1987
REAM, W.: "Agrobacterium", 2008, SPRINGER SCIENCE+BUSINESS MEDIA, LLC, article "Production of a mobile T-DNA by Agrobacterium tumefaciens", pages: 280 - 313
TAO, Q.; ZHANG, H.-B.: "Cloning and stable maintenance of DNA fragments over 300 kb in Escherichia coli with conventional plasmid-based vectors", NUCLEIC ACIDS RESEARCH, vol. 26, 1998, pages 4901 - 4909, XP002933437, DOI: doi:10.1093/nar/26.21.4901
AMOAH, B.K.; WU, H.; SPARKS, C.; JONES, H.D.: "Factors influencing Agrobacterium-mediated transient expression of uidA in wheat inflorescence tissue", J EXP BOT, vol. 52, 2001, pages 1135 - 1142, XP002995060, DOI: doi:10.1093/jexbot/52.358.1135
AN, G.; EVERT, P.R.; MITRA, A.; HA, S.B.: "Plant Molecular Biology Manual A3", 1988, KLUWER ACADEMIC PRESS, article "Binary vectors", pages: 1 - 19
AROKIARAJ, P.; LEELAWATHY, R; YEANG, H.Y.: "The supervirulence plasmid pTOK47 from Agrobacterium tumefaciens A281 improves transformation efficiency of Hevea brasiliensis", AM J BIOCHEM BIOTECHNOL, vol. 5, 2009, pages 137 - 141
BEVAN, M.: "Binary Agrobacterium vectors for plant transformation", NUCLEIC ACIDS RES, vol. 12, 1984, pages 8711 - 8721
CHRISTIE, P. J.: "Type IV secretion: the Agrobacterium VirB/D4 and related conjugation systems", BIOCHIM BIOPYS ACTA, vol. 1694, 2004, pages 219 - 234, XP004687667, DOI: doi:10.1016/j.bbamcr.2004.02.013
CITOVSKY, V.; DE VOS, G.; ZAMBRYSKI, P.: "Single-stranded DNA binding protein encoded by the virE locus of Agrobacterium tumefaciens", SCIENCE, vol. 240, 1988, pages 501 - 504
CLOSE, T.J.; TAIT, R.C.; REMPEL, H.C.; HIROOKA, T.; KIM, L.; KADO, C.I.: "Molecular characterization of the virC genes of the Ti plasmid", J BACTERIOL, vol. 169, 1987, pages 2336 - 2344
DITTA, G.; STANFIELD, S.; CORBIN, D.; HELINSKI, D.R.: "Broad host range DNA cloning system for gram-negative bacteria: construction of gene bank of Rhizobium meliloti.", PROC NATL ACAD SCI USA, vol. 77, 1980, pages 7347 - 7351
DONG, S.; QU, R.: "High efficiency transformation of tall fescue with Agrobacterium tumefaciens", PLANT SCI, vol. 168, 2005, pages 1453 - 1458, XP025296148, DOI: doi:10.1016/j.plantsci.2005.01.008
FRALEY, RT.; ROGERS, S.G.; HORSCH, RB.; EICHOLTZ, D.A.; FLICK, J.S.: "The SEV system: a new disarmed Ti plasmid vector for plant transformation", BIOTECHNOLOGY, vol. 3, 1985, pages 629 - 635
FRALEY, RT.; ROGERS, S.G.; HORSCH, RB.; SANDERS, P.R.; FLICK, J.S.; ADAMS, S.P.; BITTNER, M.L.; BRAND, L.A.; FINK, C.L.; FRY, J.S.: "Expression of bacterial genes in plant cells", PROC NATL ACAD SCI USA, vol. 80, 1983, pages 4803 - 4807, XP001062995, DOI: doi:10.1073/pnas.80.15.4803
HAMILTON, C.M.; FRARY, A.; LEWIS, C.; TANKSLEY, S.D.: "Stable transfer of intact high molecular weight DNA into plant chromosomes", PROC NATL ACAD SCI USA, vol. 93, 1996, pages 9975 - 9979, XP002263681, DOI: doi:10.1073/pnas.93.18.9975
HELLENS, R.P.; EDWARDS, E.A.; LEYLAND, N.R.; BEAN, S.; MULLINEAUX, P.M.: "pGreen: a versatile and flexible binary Ti vector for Agrobacterium-mediated plant transformation", PLANT MOL BIOL, vol. 42, 2000, pages 819 - 832, XP002959415, DOI: doi:10.1023/A:1006496308160
HEPBURN, A.G.; WHITE, J.; PEARSON, L.; MAUNDERS, M.J.; CLARKE, L.E.; PRESCOTT, A.G.; BLUNDY, K.S.: "The use of pNJ5000 as an intermediate vector for the genetic manipulation of Agrobacterium Ti-plasmids", J GEN MICROBIOL, vol. 131, 1985, pages 2961 - 2969
HIEI, Y.; KOMARI, T.: "Improved protocols for transformation of indica rice mediated by Agrobacterium tumefaciens", PLANT CELL TISSUE ORGAN CULT, vol. 85, 2006, pages 271 - 283, XP019409331, DOI: doi:10.1007/s11240-005-9069-8
HIEI, Y.; KOMARI, T.: "Agrobacterium-mediated transformation of rice using immature embryos or calli induced from mature seed", NAT PROTOCOLS, vol. 3, 2008, pages 824 - 834
HIEI, Y.; OHTA, S.; KOMARI, T.; KUMASHIRO, T.: "Efficient transformation of rice (Oryza sativa L.) mediated by Agrobacterium and sequence analysis of the boundaries of the T-DNA", PLANT J, vol. 6, 1994, pages 271 - 282
HOEKEMA, A.; HIRSCH, P.R.; HOOYKAAS, P.J.J.; SCHILPEROORT, R.A.: "A binary plant vector strategy based on separation of vir- and T-region of the Agrobacterium tumefaciens Ti-plasmid", NATURE, vol. 303, 1983, pages 179 - 180, XP000941788, DOI: doi:10.1038/303179a0
HOOD, E.E.; HELMER, G.L.; FRALEY, RT.; CHILTON, M.-D.: "The hypervirulence of Agrobacterium tumefaciens A281 is encoded in a region of pTiBo542 outside of T-DNA", J BACTERIOL, vol. 168, 1986, pages 1291 - 1301, XP002676672
HOOD, E.E.; GELVIN, S.B.; MELCHERS, L.S.; HOEKEMA, A.: "New Agrobacterium helper plasmids for gene transfer to plants", TRANSGENIC RES, vol. 2, 1993, pages 208 - 218, XP000197154, DOI: doi:10.1007/BF01977351
ISHIDA, Y.; HIEI, Y.; KOMARI, T.: "Agrobacterium-mediated transformation of maize", NAT PROTOCOLS, vol. 2, 2007, pages 1614 - 1621, XP009148413, DOI: doi:10.1038/nprot.2007.241
ISHIDA, Y.; SAITO, H.; OHTA, S.; HIEI, Y.; KOMARI, T.; KUMASHIRO, T: "High efficiency transformation of maize (Zea mays L.) mediated by Agrobacterium tumefaciens", NAT BIOTECHNOL, vol. 14, 1996, pages 745 - 750, XP002046364, DOI: doi:10.1038/nbt0696-745
JEFFERSON, R.A.: "Assaying chimeric genes in plants: the GUS gene fusion system", PLANT MOL. BIOL. REP., vol. 5, 1987, pages 387 - 405, XP002906650
JOUANIN, L.; GUERCHE, P.; PAMBOUKDJIAN, N.; TOURNEUR, C.; DELBART, F.C.; TOURNEUR, J.: "Structure of T-DNA in plants regenerated from roots transformed by Agrobacterium rhizogenes strain A4", MOL GEN GENET, vol. 206, 1987, pages 387 - 392
KHANNA, H.K.; DAGGARD, G.D.: "Agrobacterium tumefaciens-mediated transformation of wheat using a superbinary vector and a polyamine-supplemented regeneration medium", PLANT CELL REP, vol. 5, 2003, pages 429 - 436, XP002982472
KOMARI, T.: "Genetic characterization of double-flowered tobacco plant obtained in a transformation experiment", THEOR APPL GENET, vol. 80, 1990, pages 167 - 171
KOMARI, T.; HALPERIN, W.; NESTER, E.W: "Physical and functional map of supervirulent Agrobacterium tumefaciens tumor-inducing plasmid pTiBo542", J BACTERIOL, vol. 166, 1986, pages 88 - 94, XP000918204
KOMARI, T.; HIEI, Y.; SAITO, Y.; MURAI, N.; KUMASHIRO, T.: "Vectors carrying two separate T-DNAs for co-transformation of higher plants mediated by Agrobacterium tumefaciens and segregation of transformants free from selection markers", PLANT J, vol. 10, 1996, pages 165 - 174
KONCZ, C.; SCHELL, J.: "The promoter of TL-DNA gene 5 controls the tissue-specific expression of chimeric genes carried by a novel type of Agrobacterium binary vector", MOL GEN GENET, vol. 204, 1986, pages 383 - 396
LAZO, G.R.; STEIN, P.A.; LUDWIG, R.A.: "A DNA transformation-competent Arabidopsis genomic library", AGROBACTERIUM BIOTECHNOLOGY, vol. 9, 1991, pages 963 - 967, XP008062132, DOI: doi:10.1038/nbt1091-963
LEE, L.-Y.; GELVIN, S.B.: "T-DNA binary vectors and systems", PLANT PHYSIOL, vol. 146, 2008, pages 325 - 332, XP002487764, DOI: doi:10.1104/pp.107.113001
MCBRIDE, K.E.; SUMMERFELT, K.R.: "Improved binary vectors for Agrobacterium-mediated plant transformation", PLANT MOL BIOL, vol. 14, 1990, pages 269 - 276, XP002000712, DOI: doi:10.1007/BF00018567
OKUMURA; KADO: "The region essential for efficient autonomous replication of pSa in Escherichia coli", MOL GEN GENET, vol. 235, 1992, pages 55 - 63, XP003024413, DOI: doi:10.1007/BF00286181
PALANICHELVAM, K.; OGER, P.; CLOUGH, S.J.; CHA, C.; BENT, A.F.; FARRAND, S.K: "A second T-region of the soybean-supervirulent chrysopine-type Ti plasmid pTiChry5, and construction of a fully disarmed vir helper plasmid", MOL PLANT MICROBE INTERACT, vol. 13, 2000, pages 1081 - 1091, XP002676671, DOI: doi:10.1094/MPMI.2000.13.10.1081
PANTOJA, M.; CHEN, L.; CHEN, Y.; NESTER, E.W.: "Agrobacterium type IV secretion is a two-step process in which export substrates associate with the virulence protein VirJ in the periplasm", MOL MICROBIOL, vol. 45, 2002, pages 1325 - 1335
BILANG, R; FUTTERER, J.; SAUTTER, C.; SCHROTTE, M.; SPANGENBERG, G.: "Agricultural Biotechnology", 1997, MERCEL DEKKER INC., article "Genetic engineering of crop plants", pages: 119 - 159
SAITO, Y.; KOMARI, T.; MASUTA, C.; HAYASHI, Y.; KUMASHIRO, T.; TAKANAMI, Y: "Cucumber mosaic virus-tolerant transgenic tomato plants expressing a satellite RNA", THEOR APPL GENET, vol. 83, 1992, pages 679 - 683
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HIEI Y. ET AL.: "Efficient transformation of rice (Oryza sativa L.) mediated by Agrobacterium and sequence analysis of the boundaries of the T-DNA", PLANT J., vol. 6, no. 2, August 1994 (1994-08-01), pages 271 - 282, XP002922710
AMOAH B.K. ET AL.: "Factors influencing Agrobacterium-mediated transient expression of uidA in wheat inflorescence tissue", J. EXP. BOT., vol. 52, no. 358, May 2001 (2001-05-01), pages 1135 - 1142, XP002995060
JIN, S.; KOMARI, T.; GORDON, M.P.; NESTER, E.W.: "Genes responsible for the supervirulence phenotype of Agrobacterium tumefaciens A281", J. BACTERIOL., vol. 169, 1987, pages 4417 - 4425, XP008101731, DOI: doi:10.1128/jb.169.10.4417-4425.1987
REAM, W.: "Agrobacterium", 2008, SPRINGER SCIENCE+BUSINESS MEDIA, LLC, article "Production of a mobile T-DNA by Agrobacterium tumefaciens", pages: 280 - 313
TAO, Q.; ZHANG, H.-B.: "Cloning and stable maintenance of DNA fragments over 300 kb in Escherichia coli with conventional plasmid-based vectors", NUCLEIC ACIDS RESEARCH, vol. 26, 1998, pages 4901 - 4909, XP002933437, DOI: doi:10.1093/nar/26.21.4901
AMOAH, B.K.; WU, H.; SPARKS, C.; JONES, H.D.: "Factors influencing Agrobacterium-mediated transient expression of uidA in wheat inflorescence tissue", J EXP BOT, vol. 52, 2001, pages 1135 - 1142, XP002995060, DOI: doi:10.1093/jexbot/52.358.1135
AN, G.; EVERT, P.R.; MITRA, A.; HA, S.B.: "Plant Molecular Biology Manual A3", 1988, KLUWER ACADEMIC PRESS, article "Binary vectors", pages: 1 - 19
AROKIARAJ, P.; LEELAWATHY, R; YEANG, H.Y.: "The supervirulence plasmid pTOK47 from Agrobacterium tumefaciens A281 improves transformation efficiency of Hevea brasiliensis", AM J BIOCHEM BIOTECHNOL, vol. 5, 2009, pages 137 - 141
BEVAN, M.: "Binary Agrobacterium vectors for plant transformation", NUCLEIC ACIDS RES, vol. 12, 1984, pages 8711 - 8721
CHRISTIE, P. J.: "Type IV secretion: the Agrobacterium VirB/D4 and related conjugation systems", BIOCHIM BIOPYS ACTA, vol. 1694, 2004, pages 219 - 234, XP004687667, DOI: doi:10.1016/j.bbamcr.2004.02.013
CITOVSKY, V.; DE VOS, G.; ZAMBRYSKI, P.: "Single-stranded DNA binding protein encoded by the virE locus of Agrobacterium tumefaciens", SCIENCE, vol. 240, 1988, pages 501 - 504
CLOSE, T.J.; TAIT, R.C.; REMPEL, H.C.; HIROOKA, T.; KIM, L.; KADO, C.I.: "Molecular characterization of the virC genes of the Ti plasmid", J BACTERIOL, vol. 169, 1987, pages 2336 - 2344
DITTA, G.; STANFIELD, S.; CORBIN, D.; HELINSKI, D.R.: "Broad host range DNA cloning system for gram-negative bacteria: construction of gene bank of Rhizobium meliloti.", PROC NATL ACAD SCI USA, vol. 77, 1980, pages 7347 - 7351
DONG, S.; QU, R.: "High efficiency transformation of tall fescue with Agrobacterium tumefaciens", PLANT SCI, vol. 168, 2005, pages 1453 - 1458, XP025296148, DOI: doi:10.1016/j.plantsci.2005.01.008
FRALEY, RT.; ROGERS, S.G.; HORSCH, RB.; EICHOLTZ, D.A.; FLICK, J.S.: "The SEV system: a new disarmed Ti plasmid vector for plant transformation", BIOTECHNOLOGY, vol. 3, 1985, pages 629 - 635
FRALEY, RT.; ROGERS, S.G.; HORSCH, RB.; SANDERS, P.R.; FLICK, J.S.; ADAMS, S.P.; BITTNER, M.L.; BRAND, L.A.; FINK, C.L.; FRY, J.S.: "Expression of bacterial genes in plant cells", PROC NATL ACAD SCI USA, vol. 80, 1983, pages 4803 - 4807, XP001062995, DOI: doi:10.1073/pnas.80.15.4803
HAMILTON, C.M.; FRARY, A.; LEWIS, C.; TANKSLEY, S.D.: "Stable transfer of intact high molecular weight DNA into plant chromosomes", PROC NATL ACAD SCI USA, vol. 93, 1996, pages 9975 - 9979, XP002263681, DOI: doi:10.1073/pnas.93.18.9975
HELLENS, R.P.; EDWARDS, E.A.; LEYLAND, N.R.; BEAN, S.; MULLINEAUX, P.M.: "pGreen: a versatile and flexible binary Ti vector for Agrobacterium-mediated plant transformation", PLANT MOL BIOL, vol. 42, 2000, pages 819 - 832, XP002959415, DOI: doi:10.1023/A:1006496308160
HEPBURN, A.G.; WHITE, J.; PEARSON, L.; MAUNDERS, M.J.; CLARKE, L.E.; PRESCOTT, A.G.; BLUNDY, K.S.: "The use of pNJ5000 as an intermediate vector for the genetic manipulation of Agrobacterium Ti-plasmids", J GEN MICROBIOL, vol. 131, 1985, pages 2961 - 2969
HIEI, Y.; KOMARI, T.: "Improved protocols for transformation of indica rice mediated by Agrobacterium tumefaciens", PLANT CELL TISSUE ORGAN CULT, vol. 85, 2006, pages 271 - 283, XP019409331, DOI: doi:10.1007/s11240-005-9069-8
HIEI, Y.; KOMARI, T.: "Agrobacterium-mediated transformation of rice using immature embryos or calli induced from mature seed", NAT PROTOCOLS, vol. 3, 2008, pages 824 - 834
HIEI, Y.; OHTA, S.; KOMARI, T.; KUMASHIRO, T.: "Efficient transformation of rice (Oryza sativa L.) mediated by Agrobacterium and sequence analysis of the boundaries of the T-DNA", PLANT J, vol. 6, 1994, pages 271 - 282
HOEKEMA, A.; HIRSCH, P.R.; HOOYKAAS, P.J.J.; SCHILPEROORT, R.A.: "A binary plant vector strategy based on separation of vir- and T-region of the Agrobacterium tumefaciens Ti-plasmid", NATURE, vol. 303, 1983, pages 179 - 180, XP000941788, DOI: doi:10.1038/303179a0
HOOD, E.E.; HELMER, G.L.; FRALEY, RT.; CHILTON, M.-D.: "The hypervirulence of Agrobacterium tumefaciens A281 is encoded in a region of pTiBo542 outside of T-DNA", J BACTERIOL, vol. 168, 1986, pages 1291 - 1301, XP002676672
HOOD, E.E.; GELVIN, S.B.; MELCHERS, L.S.; HOEKEMA, A.: "New Agrobacterium helper plasmids for gene transfer to plants", TRANSGENIC RES, vol. 2, 1993, pages 208 - 218, XP000197154, DOI: doi:10.1007/BF01977351
ISHIDA, Y.; HIEI, Y.; KOMARI, T.: "Agrobacterium-mediated transformation of maize", NAT PROTOCOLS, vol. 2, 2007, pages 1614 - 1621, XP009148413, DOI: doi:10.1038/nprot.2007.241
ISHIDA, Y.; SAITO, H.; OHTA, S.; HIEI, Y.; KOMARI, T.; KUMASHIRO, T: "High efficiency transformation of maize (Zea mays L.) mediated by Agrobacterium tumefaciens", NAT BIOTECHNOL, vol. 14, 1996, pages 745 - 750, XP002046364, DOI: doi:10.1038/nbt0696-745
JEFFERSON, R.A.: "Assaying chimeric genes in plants: the GUS gene fusion system", PLANT MOL. BIOL. REP., vol. 5, 1987, pages 387 - 405, XP002906650
JOUANIN, L.; GUERCHE, P.; PAMBOUKDJIAN, N.; TOURNEUR, C.; DELBART, F.C.; TOURNEUR, J.: "Structure of T-DNA in plants regenerated from roots transformed by Agrobacterium rhizogenes strain A4", MOL GEN GENET, vol. 206, 1987, pages 387 - 392
KHANNA, H.K.; DAGGARD, G.D.: "Agrobacterium tumefaciens-mediated transformation of wheat using a superbinary vector and a polyamine-supplemented regeneration medium", PLANT CELL REP, vol. 5, 2003, pages 429 - 436, XP002982472
KOMARI, T.: "Genetic characterization of double-flowered tobacco plant obtained in a transformation experiment", THEOR APPL GENET, vol. 80, 1990, pages 167 - 171
KOMARI, T.; HALPERIN, W.; NESTER, E.W: "Physical and functional map of supervirulent Agrobacterium tumefaciens tumor-inducing plasmid pTiBo542", J BACTERIOL, vol. 166, 1986, pages 88 - 94, XP000918204
KOMARI, T.; HIEI, Y.; SAITO, Y.; MURAI, N.; KUMASHIRO, T.: "Vectors carrying two separate T-DNAs for co-transformation of higher plants mediated by Agrobacterium tumefaciens and segregation of transformants free from selection markers", PLANT J, vol. 10, 1996, pages 165 - 174
KONCZ, C.; SCHELL, J.: "The promoter of TL-DNA gene 5 controls the tissue-specific expression of chimeric genes carried by a novel type of Agrobacterium binary vector", MOL GEN GENET, vol. 204, 1986, pages 383 - 396
LAZO, G.R.; STEIN, P.A.; LUDWIG, R.A.: "A DNA transformation-competent Arabidopsis genomic library", AGROBACTERIUM BIOTECHNOLOGY, vol. 9, 1991, pages 963 - 967, XP008062132, DOI: doi:10.1038/nbt1091-963
LEE, L.-Y.; GELVIN, S.B.: "T-DNA binary vectors and systems", PLANT PHYSIOL, vol. 146, 2008, pages 325 - 332, XP002487764, DOI: doi:10.1104/pp.107.113001
MCBRIDE, K.E.; SUMMERFELT, K.R.: "Improved binary vectors for Agrobacterium-mediated plant transformation", PLANT MOL BIOL, vol. 14, 1990, pages 269 - 276, XP002000712, DOI: doi:10.1007/BF00018567
OKUMURA; KADO: "The region essential for efficient autonomous replication of pSa in Escherichia coli", MOL GEN GENET, vol. 235, 1992, pages 55 - 63, XP003024413, DOI: doi:10.1007/BF00286181
PALANICHELVAM, K.; OGER, P.; CLOUGH, S.J.; CHA, C.; BENT, A.F.; FARRAND, S.K: "A second T-region of the soybean-supervirulent chrysopine-type Ti plasmid pTiChry5, and construction of a fully disarmed vir helper plasmid", MOL PLANT MICROBE INTERACT, vol. 13, 2000, pages 1081 - 1091, XP002676671, DOI: doi:10.1094/MPMI.2000.13.10.1081
PANTOJA, M.; CHEN, L.; CHEN, Y.; NESTER, E.W.: "Agrobacterium type IV secretion is a two-step process in which export substrates associate with the virulence protein VirJ in the periplasm", MOL MICROBIOL, vol. 45, 2002, pages 1325 - 1335
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