WO1996017945B1 - Reversible nuclear genetic system for male sterility in transgenic plants - Google Patents
Reversible nuclear genetic system for male sterility in transgenic plantsInfo
- Publication number
- WO1996017945B1 WO1996017945B1 PCT/US1995/015229 US9515229W WO9617945B1 WO 1996017945 B1 WO1996017945 B1 WO 1996017945B1 US 9515229 W US9515229 W US 9515229W WO 9617945 B1 WO9617945 B1 WO 9617945B1
- Authority
- WO
- WIPO (PCT)
- Prior art keywords
- promoter
- plant
- anther
- dna sequence
- male sterility
- Prior art date
Links
- 206010021929 Infertility male Diseases 0.000 title claims abstract 3
- 208000007466 Male Infertility Diseases 0.000 title claims abstract 3
- 230000002068 genetic effect Effects 0.000 title abstract 3
- 230000002441 reversible effect Effects 0.000 title 1
- 230000009261 transgenic effect Effects 0.000 title 1
- 108091028043 Nucleic acid sequence Proteins 0.000 claims abstract 6
- 108090000623 proteins and genes Proteins 0.000 claims abstract 6
- 230000015572 biosynthetic process Effects 0.000 claims abstract 3
- 101710112752 Cytotoxin Proteins 0.000 claims 1
- 102000053602 DNA Human genes 0.000 claims 1
- 108010031746 Dam methyltransferase Proteins 0.000 claims 1
- 101000653753 Escherichia phage Mu Tail fiber protein S Proteins 0.000 claims 1
- 108020004511 Recombinant DNA Proteins 0.000 claims 1
- 231100000599 cytotoxic agent Toxicity 0.000 claims 1
- 239000002619 cytotoxin Substances 0.000 claims 1
- 238000010348 incorporation Methods 0.000 abstract 1
- 230000008121 plant development Effects 0.000 abstract 1
- 230000001105 regulatory effect Effects 0.000 abstract 1
- 238000009877 rendering Methods 0.000 abstract 1
- 230000001131 transforming effect Effects 0.000 abstract 1
Abstract
Plant development can be altered by transforming a plant with a genetic construct that includes regulatory elements and DNA sequences capable of acting in a fashion to inhibit pollen formation or function, thus rendering the transformed plant reversibly male-sterile. In particular, the present invention relates to the use of dominant negative genes and an anther-specific promoter. Male sterility is reversed by incorporation into a plant of a second genetic construct which represses the dominant negative gene. The invention also relates to novel DNA sequences which exhibit the ability to serve as anther-specific promoters in plants.
Claims
AMENDED CLAIMS
[received by the International Bureau on 18 June 1996 (18.06.96); new claims 29 - 38 added; remaining claims unchanged (1 page)]
29. A method of producing a male sterile plant comprising the steps of : producing a transformed plant comprising a recombinant DNA molecule which comprises (i) a DNA sequence encoding a gene product which inhibits pollen formation or function and (ii) a promoter specific to cells critical to pollen formation or function operatively linked to said DNA sequence encoding said gene product; and growing said plant under conditions such that male sterility is achieved as a result of the expression of said DNA sequence .
30. The method of claim 29 wherein said gene product s a cytotoxin. 31. The method of claim 29 wherein said promoter is an anther-specific promoter.
32. The method of claim 31 wherein said anther- specific promoter is 5126 promoter.
33. The method of claim 29, wherein said DNA sequence encodes an exogenous methylase gene.
34. The method of claim 33, wherein said exogenous methylase gene is the DAM methylase gene.
35. The method of claim 34, wherein said promoter is an anther-specific promoter. 36. The method of claim 35, wherein said anther- specific promoter is 5126 promoter.
37. A plant produced by the method of claim 29.
38. A plant produced by the method of claim 1.
Priority Applications (10)
| Application Number | Priority Date | Filing Date | Title |
|---|---|---|---|
| NZ297303A NZ297303A (en) | 1994-12-08 | 1995-12-07 | Producing reversible male sterility in plants using a reversible nuclear genetic system |
| BR9510456A BR9510456A (en) | 1994-12-08 | 1995-12-07 | Process for the production of reversible male sterility in a plant |
| CA002207264A CA2207264C (en) | 1994-12-08 | 1995-12-07 | Reversible nuclear genetic system for male sterility in transgenic plants |
| DE69535040T DE69535040T2 (en) | 1994-12-08 | 1995-12-07 | REVERSIBLE CORE GENETIC SYSTEM FOR MALE STERILITY IN TRANSGENIC PLANTS |
| EP95940806A EP0797675B1 (en) | 1994-12-08 | 1995-12-07 | Reversible nuclear genetic system for male sterility in transgenic plants |
| JP51762796A JP3370677B2 (en) | 1994-12-08 | 1995-12-07 | A reversible nuclear genetic system for male sterility in transgenic plants |
| KR1019970703834A KR100278819B1 (en) | 1994-12-08 | 1995-12-07 | Reversible nuclear genetic system for male sterility in transgenic plants |
| MX9704270A MX9704270A (en) | 1994-12-08 | 1995-12-07 | Reversible nuclear genetic system for male sterility in transgenic plants. |
| HU9701839A HU226920B1 (en) | 1994-12-08 | 1995-12-07 | Reversible nuclear genetic system for male sterility in transgenic plants |
| AU42436/96A AU698286B2 (en) | 1994-12-08 | 1995-12-07 | Reversible nuclear genetic system for male sterility in transgenic plants |
Applications Claiming Priority (4)
| Application Number | Priority Date | Filing Date | Title |
|---|---|---|---|
| US08/351,899 | 1994-12-08 | ||
| US08/351,899 US5750868A (en) | 1994-12-08 | 1994-12-08 | Reversible nuclear genetic system for male sterility in transgenic plants |
| US08/474,556 US5689051A (en) | 1994-12-08 | 1995-06-07 | Transgenic plants and DNA comprising anther specific promoter 5126 and gene to achieve male sterility |
| US08/474,556 | 1995-06-07 |
Publications (2)
| Publication Number | Publication Date |
|---|---|
| WO1996017945A1 WO1996017945A1 (en) | 1996-06-13 |
| WO1996017945B1 true WO1996017945B1 (en) | 1996-07-25 |
Family
ID=26997302
Family Applications (1)
| Application Number | Title | Priority Date | Filing Date |
|---|---|---|---|
| PCT/US1995/015229 WO1996017945A1 (en) | 1994-12-08 | 1995-12-07 | Reversible nuclear genetic system for male sterility in transgenic plants |
Country Status (13)
| Country | Link |
|---|---|
| US (5) | US5837851A (en) |
| EP (3) | EP0797675B1 (en) |
| JP (2) | JP3370677B2 (en) |
| AR (1) | AR000291A1 (en) |
| AT (1) | ATE329040T1 (en) |
| AU (1) | AU698286B2 (en) |
| DE (1) | DE69535040T2 (en) |
| ES (1) | ES2267105T3 (en) |
| HU (2) | HU226920B1 (en) |
| MX (1) | MX9704270A (en) |
| NZ (1) | NZ297303A (en) |
| PT (1) | PT797675E (en) |
| WO (1) | WO1996017945A1 (en) |
Families Citing this family (59)
| Publication number | Priority date | Publication date | Assignee | Title |
|---|---|---|---|---|
| GB8901677D0 (en) | 1989-01-26 | 1989-03-15 | Ici Plc | Hybrid seed production |
| US5837850A (en) * | 1994-04-21 | 1998-11-17 | Pioneer Hi-Bred International, Inc. | Regulatory element conferring tapetum specificity |
| US5837851A (en) * | 1994-12-08 | 1998-11-17 | Pioneer Hi-Bred International, Inc. | DNA promoter 5126 and constructs useful in a reversible nuclear genetic system for male sterility in transgenic plants |
| US5750868A (en) * | 1994-12-08 | 1998-05-12 | Pioneer Hi-Bred International, Inc. | Reversible nuclear genetic system for male sterility in transgenic plants |
| US5962769A (en) * | 1995-06-07 | 1999-10-05 | Pioneer Hi-Bred International, Inc. | Induction of male sterility in plants by expression of high levels of avidin |
| US7154024B2 (en) * | 1997-06-23 | 2006-12-26 | Pioneer Hi-Bred, Inc. | Male tissue-preferred regulatory sequences of Ms45 gene and method of using same |
| US6037523A (en) * | 1997-06-23 | 2000-03-14 | Pioneer Hi-Bred International | Male tissue-preferred regulatory region and method of using same |
| JP2002504343A (en) * | 1998-02-20 | 2002-02-12 | シンジェンタ リミテッド | Hybrid seed production |
| AU3040200A (en) * | 1998-12-21 | 2000-07-12 | Max-Planck-Gesellschaft Zur Forderung Der Wissenschaften E.V. | Novel mads-box genes and uses thereof |
| JP4642239B2 (en) * | 1998-12-22 | 2011-03-02 | ダウ・アグロサイエンス・エル・エル・シー | Methods and genetic compositions for limiting outcrossing and undesirable gene spreading in crop plants |
| WO2000055315A1 (en) | 1999-03-16 | 2000-09-21 | Pioneer Hi-Bred International, Inc. | Sf3 promoter and methods of use |
| WO2000058454A1 (en) * | 1999-03-26 | 2000-10-05 | Japan As Represented By Director General Of National Institute Of Agrobiological Resources, Ministry Of Agriculture, Forestry And Fisheries | Promoter sequences expressed in anthers and pollens |
| WO2001000834A1 (en) * | 1999-06-29 | 2001-01-04 | Pioneer Hi-Bred International, Inc. | Gene affecting male fertility in plants |
| AU3817701A (en) * | 2000-02-15 | 2001-08-27 | Pioneer Hi Bred Int | Male tissue-preferred regulatory region and method of using same |
| US6838593B2 (en) * | 2000-07-13 | 2005-01-04 | Pioneer Hi-Bred Int'l Inc. | Austin responsive promoter sequences and methods of using the same |
| EP1319081A2 (en) * | 2000-09-18 | 2003-06-18 | Pioneer Hi-Bred International, Inc. | Rice mlh1 ortholog and uses thereof |
| BR0114199A (en) * | 2000-09-26 | 2006-01-31 | Pioneer Hi Bred Int | Nucleotide sequences measuring male fertility and methods of using them |
| US7517975B2 (en) | 2000-09-26 | 2009-04-14 | Pioneer Hi-Bred International, Inc. | Nucleotide sequences mediating male fertility and method of using same |
| US7612251B2 (en) | 2000-09-26 | 2009-11-03 | Pioneer Hi-Bred International, Inc. | Nucleotide sequences mediating male fertility and method of using same |
| US20020129399A1 (en) * | 2001-03-09 | 2002-09-12 | Marc Albertsen | Biotin-binding compounds for induction of sterility in plants |
| US7230168B2 (en) * | 2001-12-20 | 2007-06-12 | The Curators Of The University Of Missouri | Reversible male sterility in transgenic plants by expression of cytokinin oxidase |
| US20050257293A1 (en) * | 2002-09-17 | 2005-11-17 | Mascia Peter N | Biological containment system |
| US7476777B2 (en) * | 2002-09-17 | 2009-01-13 | Ceres, Inc. | Biological containment system |
| CA2442521A1 (en) * | 2002-10-03 | 2004-04-03 | Derek Lydiate | Repressor-mediated selection strategies |
| US20040133944A1 (en) * | 2003-01-08 | 2004-07-08 | Delta And Pine Land Company | Seed oil suppression to enhance yield of commercially important macromolecules |
| US20050044596A1 (en) * | 2003-03-19 | 2005-02-24 | Smith Alan G. | Methods to confer enhanced floral properties to plants |
| EP1616949A4 (en) * | 2003-04-07 | 2006-06-21 | Jujo Paper Co Ltd | Novel vector and method of constructing transformant plant using the vector |
| KR100537955B1 (en) | 2003-10-29 | 2005-12-20 | 학교법인고려중앙학원 | A solely pollen-specific promoter |
| EP2333075B1 (en) | 2003-12-16 | 2018-07-04 | Pioneer Hi-Bred International Inc. | Dominant gene suppression transgenes and methods of using same |
| US20070169227A1 (en) | 2003-12-16 | 2007-07-19 | Pioneer Hi-Bred International Inc. | Dominant Gene Suppression Transgenes and Methods of Using Same |
| WO2006080598A1 (en) * | 2005-01-31 | 2006-08-03 | Dongbu Hitek Co., Ltd. | Anther-specific genes derived from malus x domestica, promoters thereof and a method for preparing transformed plant using the same |
| PL1863334T3 (en) | 2005-03-21 | 2012-04-30 | Univ California | Methods to generate drought-resistant plants |
| US7550579B2 (en) | 2005-04-29 | 2009-06-23 | Pioneer Hi-Bred International, Inc. | Pericarp-preferred regulatory element |
| WO2006125678A2 (en) * | 2005-05-23 | 2006-11-30 | Nunhems Netherlands Bv | Reversible male-sterile plants |
| US7910802B2 (en) | 2007-08-03 | 2011-03-22 | Pioneer Hi-Bred International, Inc. | MSCA1 nucleotide sequences impacting plant male fertility and method of using same |
| US7919676B2 (en) | 2007-08-03 | 2011-04-05 | Pioneer Hi-Bred International, Inc. | Msca1 nucleotide sequences impacting plant male fertility and method of using same |
| US7915478B2 (en) | 2007-08-03 | 2011-03-29 | Pioneer Hi-Bred International, Inc. | Msca1 nucleotide sequences impacting plant male fertility and method of using same |
| BRPI0722195B1 (en) | 2007-08-03 | 2018-01-30 | Pioneer Hi-Bred Internacional, Inc | METHODS FOR MAINTENANCE OF A PLANT RECESSIVE HOMOZYGOTE STATE |
| AR075465A1 (en) | 2008-10-22 | 2011-04-06 | Basf Se | USE OF SULFONILUREA HERBICIDES IN CULTIVATED PLANTS |
| WO2010046422A2 (en) | 2008-10-22 | 2010-04-29 | Basf Se | Use of auxin type herbicides on cultivated plants |
| CN104080914A (en) | 2011-06-21 | 2014-10-01 | 先锋国际良种公司 | Methods and compositions for producing male sterile plants |
| MX2014011038A (en) | 2012-03-13 | 2015-06-02 | Pioneer Hi Bred Int | Genetic reduction of male fertility in plants. |
| WO2013138309A1 (en) | 2012-03-13 | 2013-09-19 | Pioneer Hi-Bred International, Inc. | Genetic reduction of male fertility in plants |
| AR093243A1 (en) | 2012-10-01 | 2015-05-27 | Basf Se | USE OF N-TIO-ANTRANILAMIDE COMPOUNDS IN CULTIVATED PLANTS |
| WO2014079820A1 (en) | 2012-11-22 | 2014-05-30 | Basf Se | Use of anthranilamide compounds for reducing insect-vectored viral infections |
| US9803214B2 (en) | 2013-03-12 | 2017-10-31 | Pioneer Hi-Bred International, Inc. | Breeding pair of wheat plants comprising an MS45 promoter inverted repeat that confers male sterility and a construct that restores fertility |
| CN105143248A (en) | 2013-03-14 | 2015-12-09 | 先锋国际良种公司 | Maize stress related transcription factor 18 and uses thereof |
| CN105143454A (en) | 2013-03-15 | 2015-12-09 | 先锋国际良种公司 | Compositions and methods of use of ACC oxidase polynucleotides and polypeptides |
| CN105579583B (en) | 2014-01-02 | 2017-12-12 | 北京思创达科技有限公司 | The breeding of Female sterile clone and cross-breeding technology |
| EP3028573A1 (en) | 2014-12-05 | 2016-06-08 | Basf Se | Use of a triazole fungicide on transgenic plants |
| WO2016091674A1 (en) | 2014-12-12 | 2016-06-16 | Basf Se | Use of cyclaniliprole on cultivated plants |
| CA2971425A1 (en) | 2014-12-16 | 2016-06-23 | Pioneer Hi-Bred International, Inc. | Restoration of male fertility in wheat |
| BR112017021450B1 (en) | 2015-04-07 | 2021-12-28 | Basf Agrochemical Products B.V. | PEST CONTROL METHODS, PLANT HEALTH IMPROVEMENT METHOD AND COATED SEED |
| JP2018512878A (en) | 2015-04-20 | 2018-05-24 | ネオゲノミクス ラボラトリーズ， インコーポレイテッド | Methods to increase the sensitivity of next-generation sequencing |
| WO2017004599A1 (en) * | 2015-07-01 | 2017-01-05 | Neogenomics Laboratories, Inc. | Method for high sensitivity detection of myd88 mutations |
| WO2017031230A1 (en) | 2015-08-17 | 2017-02-23 | Neogenomics Laboratories, Inc. | High-sensitivity sequencing to detect btk inhibitor resistance |
| DE102016106656A1 (en) | 2016-04-12 | 2017-10-12 | Kws Saat Se | Nuclear encoded male sterility by mutation in cytochrome P450 oxidase |
| DE102016015741A1 (en) | 2016-04-12 | 2017-11-30 | Kws Saat Se | Nuclear encoded male sterility by mutation in cytochrome P450 oxidase |
| EP3338552A1 (en) | 2016-12-21 | 2018-06-27 | Basf Se | Use of a tetrazolinone fungicide on transgenic plants |
Family Cites Families (17)
| Publication number | Priority date | Publication date | Assignee | Title |
|---|---|---|---|---|
| US4407956A (en) * | 1981-03-13 | 1983-10-04 | The Regents Of The University Of California | Cloned cauliflower mosaic virus DNA as a plant vehicle |
| CA1192510A (en) * | 1981-05-27 | 1985-08-27 | Lawrence E. Pelcher | Rna plant virus vector or portion thereof, a method of construction thereof, and a method of producing a gene derived product therefrom |
| NL8200523A (en) * | 1982-02-11 | 1983-09-01 | Univ Leiden | METHOD FOR TRANSFORMING IN VITRO PLANT PROTOPLASTS WITH PLASMIDE DNA. |
| US4536475A (en) * | 1982-10-05 | 1985-08-20 | Phytogen | Plant vector |
| EP0116718B2 (en) * | 1983-01-13 | 1996-05-08 | Max-Planck-Gesellschaft zur Förderung der Wissenschaften e.V. | Process for the introduction of expressible genes into plant cell genomes and agrobacterium strains carrying hybrid Ti plasmid vectors useful for this process |
| EP0160692A1 (en) * | 1983-11-03 | 1985-11-13 | DE WET, Johannes Martenis Jacob | Method for the transfer of exogenous genes in plants using pollen as a vector |
| NZ219531A (en) * | 1986-03-14 | 1991-10-25 | Exxon Research Engineering Co | Anti-fouling fuel compositions and additives containing amines and amine oxides |
| EP0265556A1 (en) * | 1986-10-31 | 1988-05-04 | Max-Planck-Gesellschaft zur Förderung der Wissenschaften e.V. | Stable binary agrobacterium vectors and their use |
| EP0275069A3 (en) * | 1987-01-13 | 1990-04-25 | DNA PLANT TECHNOLOGY CORPORATION (under the laws of the state of Delaware) | Pollen-mediated gene transformation in plants |
| GB8901677D0 (en) * | 1989-01-26 | 1989-03-15 | Ici Plc | Hybrid seed production |
| DE68918494T2 (en) * | 1988-05-17 | 1995-03-23 | Lubrizol Genetics Inc | Herbal ubiquitin promoter system. |
| IL99413A0 (en) * | 1990-09-06 | 1992-08-18 | Du Pont | Compounds and constructs for producing male sterile plants |
| US5409823A (en) * | 1992-09-24 | 1995-04-25 | Ciba-Geigy Corporation | Methods for the production of hybrid seed |
| WO1994009143A1 (en) * | 1992-10-15 | 1994-04-28 | Mogen International N.V. | Genetic moderation or restoration of plant phenotypes |
| US5750868A (en) * | 1994-12-08 | 1998-05-12 | Pioneer Hi-Bred International, Inc. | Reversible nuclear genetic system for male sterility in transgenic plants |
| US5837851A (en) * | 1994-12-08 | 1998-11-17 | Pioneer Hi-Bred International, Inc. | DNA promoter 5126 and constructs useful in a reversible nuclear genetic system for male sterility in transgenic plants |
| US7848465B2 (en) | 2007-04-02 | 2010-12-07 | Broadcom Corporation | Joint decoding of ISI (inter-symbol interference) channel and modulation codes |
-
1995
- 1995-06-07 US US08/479,041 patent/US5837851A/en not_active Expired - Lifetime
- 1995-06-07 US US08/479,383 patent/US5795753A/en not_active Expired - Lifetime
- 1995-06-07 US US08/479,382 patent/US5763243A/en not_active Expired - Lifetime
- 1995-12-06 AR AR33452095A patent/AR000291A1/en unknown
- 1995-12-07 AU AU42436/96A patent/AU698286B2/en not_active Expired
- 1995-12-07 DE DE69535040T patent/DE69535040T2/en not_active Expired - Lifetime
- 1995-12-07 PT PT95940806T patent/PT797675E/en unknown
- 1995-12-07 AT AT95940806T patent/ATE329040T1/en not_active IP Right Cessation
- 1995-12-07 EP EP95940806A patent/EP0797675B1/en not_active Expired - Lifetime
- 1995-12-07 HU HU9701839A patent/HU226920B1/en not_active IP Right Cessation
- 1995-12-07 JP JP51762796A patent/JP3370677B2/en not_active Expired - Lifetime
- 1995-12-07 HU HU0301904A patent/HU227303B1/en not_active IP Right Cessation
- 1995-12-07 EP EP06011502A patent/EP1731614A1/en not_active Withdrawn
- 1995-12-07 NZ NZ297303A patent/NZ297303A/en not_active IP Right Cessation
- 1995-12-07 WO PCT/US1995/015229 patent/WO1996017945A1/en active IP Right Grant
- 1995-12-07 MX MX9704270A patent/MX9704270A/en unknown
- 1995-12-07 ES ES95940806T patent/ES2267105T3/en not_active Expired - Lifetime
- 1995-12-07 EP EP02016942A patent/EP1273662B1/en not_active Expired - Lifetime
-
1997
- 1997-03-17 US US08/819,646 patent/US6281348B1/en not_active Expired - Lifetime
-
1998
- 1998-11-19 US US09/195,716 patent/US6399856B1/en not_active Expired - Lifetime
-
2002
- 2002-03-11 JP JP2002066072A patent/JP2002354954A/en active Pending
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