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WO1993013649A1 - Plantes de ble a modification genetique, lignee obtenue de ces plantes, et procede de production de ble hybride - Google Patents

Plantes de ble a modification genetique, lignee obtenue de ces plantes, et procede de production de ble hybride Download PDF

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Publication number
WO1993013649A1
WO1993013649A1 PCT/AU1993/000017 AU9300017W WO9313649A1 WO 1993013649 A1 WO1993013649 A1 WO 1993013649A1 AU 9300017 W AU9300017 W AU 9300017W WO 9313649 A1 WO9313649 A1 WO 9313649A1
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Prior art keywords
chromosome
gene
wheat
pairing
plant
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PCT/AU1993/000017
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English (en)
Inventor
Geoffrey Frank Smart
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Pacific Seeds Pty Ltd
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Pacific Seeds Pty Ltd
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Priority to AU33388/93A priority Critical patent/AU654762B2/en
Publication of WO1993013649A1 publication Critical patent/WO1993013649A1/fr
Anticipated expiration legal-status Critical
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    • A—HUMAN NECESSITIES
    • A01—AGRICULTURE; FORESTRY; ANIMAL HUSBANDRY; HUNTING; TRAPPING; FISHING
    • A01H—NEW PLANTS OR NON-TRANSGENIC PROCESSES FOR OBTAINING THEM; PLANT REPRODUCTION BY TISSUE CULTURE TECHNIQUES
    • A01H1/00—Processes for modifying genotypes ; Plants characterised by associated natural traits
    • A01H1/02—Methods or apparatus for hybridisation; Artificial pollination ; Fertility
    • A01H1/022—Genic fertility modification, e.g. apomixis
    • A01H1/023—Male sterility
    • A—HUMAN NECESSITIES
    • A01—AGRICULTURE; FORESTRY; ANIMAL HUSBANDRY; HUNTING; TRAPPING; FISHING
    • A01H—NEW PLANTS OR NON-TRANSGENIC PROCESSES FOR OBTAINING THEM; PLANT REPRODUCTION BY TISSUE CULTURE TECHNIQUES
    • A01H5/00—Angiosperms, i.e. flowering plants, characterised by their plant parts; Angiosperms characterised otherwise than by their botanic taxonomy
    • A01H5/10—Seeds
    • A—HUMAN NECESSITIES
    • A01—AGRICULTURE; FORESTRY; ANIMAL HUSBANDRY; HUNTING; TRAPPING; FISHING
    • A01H—NEW PLANTS OR NON-TRANSGENIC PROCESSES FOR OBTAINING THEM; PLANT REPRODUCTION BY TISSUE CULTURE TECHNIQUES
    • A01H6/00—Angiosperms, i.e. flowering plants, characterised by their botanic taxonomy
    • A01H6/46—Gramineae or Poaceae, e.g. ryegrass, rice, wheat or maize
    • A01H6/4678—Triticum sp. [wheat]

Definitions

  • the present invention is directed generally to genetically modified wheat plants and to progeny arising from these plants, and also to a method of producing hybrid wheat using these plants.
  • the genetically altered wheat plant utilize homoeologous pairing to add a fertility restorer chromosome which contains telomeric chromatin which is homologous to another wheat chromosome which bears a recessive male sterility gene.
  • the fertility restorer chromosome contains a major portion of alien chromatin from other wheat species, and also contains a male fertility restorer gene, and one or more marker genes. These plants possess meioti ⁇ stability, and are used to produce hybrid wheat strains.
  • hybrid crops carrying selected desirable characteristics is of enormous commercial and economical importance to both individual farmers and to the agricultural industry as a whole.
  • One way of potentially obtaining hybrid crops expressing both "X” and “Y” characteristics (“XY” ) is to grow rows of "X” plants interspersed by rows of "Y” plants and attempt to ensure that only pollen from "X” plants pollinated “Y” plants, or vice versa. Such a procedure does not permit the production of "XY” hybrids with any degree of certainty due mainly to the bisexual nature of crop plants which generally favours self-pollination.
  • Male sterility may also be induced by treatment of the intended female parent plants with a chemical hybridising agent (CHA) which inhibits synthesis of viable pollen.
  • CHA chemical hybridising agent
  • CHA treatment is somewhat inefficient and a certain amount of self-pollination still occurs.
  • the seed hybrids which are produced using CHAs therefore tend to be contaminated with seed of the female parent with separation of the contaminant, if not impossible, being very costly.
  • CMS cytoplasmic male sterility
  • nuclear male sterility occasioned by the presence in plant cell nuclei of a gene directing expression of the sterility trait.
  • the nuclear male sterility genes are generally of the recessive type, referred to as " s".
  • the presence of the normal dominant male fertility gene is referred to as "+Ms”. Accordingly, the possible genotypes of a male fertile plant are +Ms/+Ms and +Ms/ms whereas a male sterile plant can only have a ms/ms genotype.
  • PCT/AU91/00319 describes a genetically modified plant for use in this method which is created using spontaneous or induced translocations between homologous chromosomes. More precisely these plant were modified by means of centric fusion or translocated chromosome which combined a male fertility gene and a colour marker, for example, a blue aleurone marker. The translocation was produced when alien addition chromosomes or substitution lines were combined.
  • a high pairing mutant was used to induce homoeologous recombination to facilitate the production of a translocation chromosome. Furthermore, a previously uncharacterised alien chromosome has been identified carrying both a male fertility gene and a marker. - -
  • Figure 1 shows two chromosomes of the wheat in accordance with the invention.
  • Figure 2 shows a procedure by which genetically modified plants may be produced.
  • Figure 3 shows another procedure by which genetically modified plants may be produced.
  • Figure 4 shows another procedure by which genetically modified plants may be produced.
  • Figure 5 shows another procedure by which genetically modified plants may be produced.
  • one aspect of the present invention provides a genetically modified plant and/or its progeny being monosomic for a wheat chromosome having a recessive male sterility gene.
  • the plant has an additional homoeologous chromosome in which homologous chromatin is present on the long arm telomere, the rest of the chromatin arising from alien wheat chromatin, said homoeologous chromosome having a dominant male fertility restorer gene and one or more selectable marker genes, and which has been created using induced homoeologous pairing.
  • the genetic makeup of the wheat plant in one preferred form can be described as:
  • Another aspect of the invention is a genetically modified precursor plant (and its progeny) for creating the previous plant which is characterised by containing a chromosome bearing a suppressor of pairing (Ph) gene which is not functioning, and which carries one or more genes for male sterility (ms), male fertility restorer genes (+Ms) or marker genes such as blue seed colouring or increased plant height.
  • a chromosome bearing a suppressor of pairing (Ph) gene which is not functioning, and which carries one or more genes for male sterility (ms), male fertility restorer genes (+Ms) or marker genes such as blue seed colouring or increased plant height.
  • a further aspect of the invention concerns the use of the genetically modified wheat plant to maintain a male sterile parental plant line, as generally described in PCT/AU91/00319. More particularly, this aspect concerns a method for the maintenance of a male sterile parental plant line for use in the production of hybrids, which comprises crossing a female parent with a male parent, the female parent being a homozygous male sterile plant, and the male parent being isogenic to the female but having a fertility maintainer chromosome bearing a dominant male fertility restorer gene and a marker gene which confers a selectable characteristic on progeny.
  • the method further includes harvesting from that cross a population of progeny consisting of a mixture of the two parental lines; and then physically separating the progeny on the basis of the presence of the marker.
  • this method is characterised by the fertility maintainer chromosome being a homoeologous chromosome in which wheat chromatin is present on the long arm telomere, the rest of the chromatin arising from alien wheat chromatin, whereby the homoeologous chromosome has a dominant male fertility restorer gene and one or more selectable marker genes, and the homoeologous chromosome having been created using induced homoeologous pairing as described in more detail hereafter.
  • the wheat be tetraploid or hexaploid wheat, especially bread wheat.
  • Any suitable wheat strain can be used as the starting wheat plant for the genetic alterations described in the invention.
  • the monosomic wheat chromosome is preferable chromosome 4B, although other chromosomes may be used instead, for example in mutant strains of wheat or in other wheats with different chromosome numbers, other chromosomes may be suitable for use in accordance with the present invention.
  • the alien wheat chromatin preferably comes from chromosomes in Agropyron elongatum, Ag. trichophorum, Triticum thaoudar or T ⁇ monococcum.
  • the Agropyron lines may contain a blue aleurone marker, and the T ⁇ thaoudar and T. monococcum lines may contain a blue marker gene, perhaps a height marker gene, and a male fertility restore gene, as generally described in PCT/AU91/00319.
  • a combination or mixture of chromatin from T ⁇ thaoudar and Ag. elongatum can also be used, if desired.
  • other wheat strains can be constructed or selected to contain the genes required in the invention.
  • the preferred marker genes are the blue aleurone gene that confers a blue colouring to seed, and a plant height gene that confers extra height to progeny plants.
  • Hexaploid wheat species normally have a chromosome number of 42, designated as chromosomes (1-7)AABBDD, tetraploid wheat species normally have a chromosome number of 28, designated (1-7)AABB, and diploid wheat species have a chromosome number usually of 14, which are chromosomes 1-7 of any of A, B or D.
  • Hexaploid or tetraploid species have evolved special genetic mechanisms to control pairing among homoeologous chromosomes, to prevent chromosome 4A pairing with chromosome 4B, instead of with 4A, its homologous chromosome, for instance.
  • Homoeologous pairing is known to be controlled by the interaction of genes on several chromosomes, and in wheat the major supressor or pairing (Phi) is located on chromosome 5B, and another suppressor of pairing (Ph2) occurs on chromosome 3D.
  • the suppressor of pairing gene (Ph) is non-functioning by being (a) mutated, so that it no longer suppresses homoeologous pairing such as with the p_h mutant phlb, or (b) by itself being suppressed, such as with strains of Ae. or (T. ) speltoides, or Ae.
  • the invention utilizes the high pairing mutation (phlb), or genes that suppress the suppressor of pairing, such as from T ⁇ speltoides, for example.
  • the selectable marker is a colour marker (such as the blue aleurone gene), or else may be a height marker, or a combination of a colour and height markers.
  • a colour marker such as the blue aleurone gene
  • the present invention in one preferred form utilizes the blue aleurone marker gene available on chromosome 4 of Agropyron elongatum, Ag. trichophoru , Triticum thaoudar or T. monococcum.
  • Agropyron elongatum is also sometimes known as Thinopyron elongatum.
  • a range of other suitable markers could also be used affecting colour, texture, size, weight or other physically identifiable characteristics. All such markers are encompassed by the present invention.
  • Some of the advantages of the instant invention concern the rapidity with which a male sterile population can be produced because cross-pollination is not desired and the advantage that no special male fertility restorer is required for the male parent in the final hybrid cross.
  • the present invention provides improved plant lines for use in producing hybrid crops, over the plant lines described in PCT/AU91/00319, which are essentially created by Robertsonian translocation or centric fusion. Homoeologous recombinants in accordance with the present invention, will be more stable. In the homozygous condition pairing and recombination will take place between wheat chromosomes and their alien homoeologues.
  • homoeologous recombination is a very rare event within the wheat genome.
  • recombination between wheat homoeologues and alien homoeologues does occur at much higher frequencies permitting the opportunity of detecting recombination between alien homoeologues carrying the blue aleurone marker(s) and the male fertility restorer gene.
  • the high pairing mutant, phlb is known to be a recessive mutant which occurs on the long arm of chromosome 5B (designated "5BL").
  • 5BL chromosome 5B
  • pairing occurs only between homologues. Therefore, at meiosis, chromosome 5B will only pair and recombine with chromosome 5B; similarly, chromosome 3A will only recombine with ⁇ hromosome 3A.
  • Wheat is a diploid species and a segmented hexaploid and when chromosomes are absent, compensation occurs.
  • chromosome 3A can pair and therefore can recombine with either chromosome 3B or 3D.
  • recombination can occur between an alien chromosome and wheat homoeologues (or possibly even between an alien chromosome and alien homoeologues).
  • chromosome when a chromosome is absent from the wheat genome, the plant is described as monosomic for that chromosome, having 41 chromosomes instead of the usual 42.
  • nullisomic i.e. carrying 40 chromosomes.
  • nullisomic plant When a nullisomic plant is self-pollinated, the progeny only carry 40 chromosomes.
  • a monosomic plant self-pollinates the progeny are of 3 types, with predictable frequencies:
  • the male parent of a cross is known to carry the phlb mutant and the female parent is monosomic for chromosome 5B, 75% of the progeny would have the opportunity to exhibit homoeologous recombination at meiosis.
  • Figure 1 shows in general form the homoeologous chromosomes in the plant according to the invention.
  • a wheat plant By using a non-functioning suppressor of pairing (Ph), such as mutant phlb on chromosome 5B, or T ⁇ speltoides which carries a suppressor of Phi, a wheat plant can be created which has a wheat 4B chromosome having a (recessive) male sterility gene (ms).
  • ms non-functioning suppressor of pairing
  • Its homoeologue of which the majority of chromatin is from an alien species, and which the 4B chromosome will pair with at meiosis, carries two marker genes, for blue seed colouring and for increased height, as well as a (dominant) male fertility restorer gene (+Ms).
  • the alien chromosome however has some native 4B chromatin on its long arm telomere, which will cause the chromosome to pair with its homoeologue 4B chromos
  • m monosomic (ie, lacking one chromosome)
  • n nucleophilicity factor
  • EXAMPLE 1 This example describes homoeologous recombination applied to the nuclear male sterile hybrid wheat system Ag. elongatum. This contains the blue aleurone marker gene on chromosome 4E. The steps in creating the desired plant are set out in Figure 2 of the drawings.
  • step 1 of Figure 2 a plant monosomic in chromosomes 4B and 5B is crossed with an addition line containing chromosome 4 of of Ag. elongatum, this being an alien chromosome the chromatin of which (except for the 4B long arm telomere) is to incorporated in the resulting wheat strain.
  • These strains are both readily available.
  • the progeny arising from this step is then crossed with a wheat plant with two doses of the phlb high pairing mutant.
  • the progeny of this cross is then crossed again with the phlb mutant, the resulting progeny ending up with two doses of phlb.
  • the plant from step 3 can be used directly in the next step, as the 5B(phlb) chromosome, while being recessive, is present in the plant as a singular chromosome, and will therefore be able to cause homoeologous recombination.
  • the wheat with either one or two ⁇ oses of the high pairing mutant is crossed with a plant being male sterile. Due to the presence of the high pairing mutant, homoeologous recombination occurs, and plants having blue seed and which are male sterile are selected, being 4B(ms) + 4E - 4B + 20" plants. These are male sterile, because while the male sterility (ms) gene is recessive, the homoeologous chromosome (4E - 4B ) lacks the dominant gene on 4B (+Ms).
  • the procedure can then continue with further crosses with plants which carry the male sterility gene (ms), for example, to get a desired product.
  • ms male sterility gene
  • This example concerns Homoeologous recombination applied to the nuclear male sterile hybrid wheat system.
  • Step 3 creates material which is homozygous for phlb. Therefore, these plants will exhibit homoeologous recombination. Thus, recombination between chromosome 4th (or 4m) and the telomeric region of chromosome 4BL is expected to occur.
  • step 4 combines, or repeats step 3 and introduces chromosome 4 from Agropyron elongatum, which carries a very effective blue aleurone marker.
  • chromosome 4 from Agropyron elongatum, which carries a very effective blue aleurone marker.
  • 4th is the 4thaoudar homoeologue from a T ⁇ thaoudar substitution line containing a blue aleurone marker, and a height marker.
  • This strain is obtained in the manner described in PCT/AU91/00319. It also has a dominant male fertility restorer gene (+Ms).
  • This strain lacks a normal 4B chromosome, and in place has an alien chromosome containing the marker genes and the +Ms gene. It is possible to utilize 4monococcum with the same characteristics in place of 4thaoudar.
  • step 1 the substitution 4thaoudar is the male parent (having the fertility restorer gene +Ms as well as the marker genes), which is readily available in seed banks, and is crossed with a strain ' monosomic in chromosomes 4B and 5B.
  • the progeny strain, m4th. n4B. m5B is selected by its marker characteristics, namely height and blue seed, and has one alien 4th chromosome, one 4B chromosome and one 5B chromosome. This chromosome configuration arises from the nature of chromosome transmission in monosomics, as described in Table 1 above.
  • step 2 the progeny from step 1 is crossed with a strain that is monosomic for chromosomes 4B and 5B, but its remaining 5B chromosome has the high pairing mutation (phlb).
  • the resulting marked progeny contain one chromosome with the high pairing mutant.
  • step 3 this is crossed with a strain that has two doses of the high pairing mutant (phlb), and which is male sterile; this strain obviously being the female parent.
  • the progeny of this cross has two doses of the (recessive) high pairing mutant phlb, and when this plant is crossed with a strain of wheat with the high pairing mutant and an extra chromosome(s) of 4 Ag. elongatum, homoeologous recombination occurs to give the desired genetically stable wheat, which can be used in hybrid production. Blue fertile seed are selected to verify the desired genes are linked.
  • This example describes an alternate method for producing the homoeologous recombined wheat plant, using the markers from T ; _ monococcum.
  • Step 3 creates material which is homozygous for phlb. Therefore, these plants will exhibit homoeologous recombination. Thus, recombination between chromosome 4m and the telomeric region of chromosome 4BL is expected to occur.
  • step 4 combines, or repeats step 3 and introduces chromosome 4 from Agropyron elongatum, which carries a very effective blue aleurone marker.
  • chromosome 4 from Agropyron elongatum, which carries a very effective blue aleurone marker.
  • 4m is 4monococcum, which is a substitution line containing a blue aleurone marker, and a height marker.
  • This strain is obtained in the manner described in PCT/AU91/00319. It also has a dominant male fertility restorer gene (+Ms).
  • This strain lacks a normal 4B chromosome, and in place has an alien chromosome containing the marker genes and the +Ms gene. It is possible to utilize 4thaoudar with the same characteristics in place of 4monococcum.
  • step 1 the substitution 4monococcum is the male parent (having the fertility restorer gene +Ms as well as the marker genes), which is readily available in seed banks, and is crossed with a strain monosomic in chromosomes 4B and 5B.
  • the progeny strain, m4m. n4B. m5B is selected by its marker characteristics, namely height and blue seed, and has one alien 4m chromosome, usually one 4B chromosome and one 5B chromosome. This chromosome configuration arises from the nature of chromosome transmission in monosomics, as described in Table 1 above.
  • step 2 the progeny from step 1 is crossed with a strain that is monosomic for chromosomes 4B and 5B, but its remaining 5B chromosome has the high pairing mutation (phlb).
  • the resulting marked progeny contain one chromosome with the high pairing mutant.
  • step 3 this is crossed with a strain that has one dose, of the high pairing mutant (phlb), and which is male sterile as the plant is monosomic for 4B(ms) so that the male sterility gene is expressed; this strain being the female parent.
  • the progeny of this cross has two doses of the (recessive) high pairing mutant phlb, and when this plant is crossed with a strain of wheat with the high pairing mutant and an extra two chromosome of 4 Ag. elongatum, homoeologous recombination occurs to give the desired genetically stable wheat, which can be used in hybrid production. Further crosses can be made to obtain a desired plant line for carrying out the procedure described in PCT/AU91/00319, if necessary, and if desired.
  • EXAMPLE 4 This involves the utilisation of an alien chromosome which appears to carry both the blue aleurone marker and male fertility gene.
  • telomeric 4BL Without the telomeric 4BL, the alien chromosome would not pair with normal 4B, which carries the male sterile allele on its short arm.
  • the presence of telomeric 4BL on the fertility restorer chromosome ensures that pairing will occur at meiosis.
  • a rod bivalent forms at meiosis as only homologous regions pair. This contrasts with other homologues within the cell which form ring bivalents.
  • both the alien chromosome and chromosome 4B are not regularly transmitted through the gametes. Unfortunately Cermeno and Zeller were unable to identify the alien chromosome. It is non-homologous with an Agropyron elongatum substitution. This suggests that the alien chromosome may be a different Agropyron species, or a diploid wheat. If the former is true then the transfer of telomeric 4BL to this alien chromosome would produce the necessary recombinant.
  • EXAMPLE 5 Although the method described in Example 4 will produce the desired recombinant alien chromosome, a more efficient method does exist. This method involves crossing the critical chromosomes into high pairing Aegilops speltoides. High pairing Aegilops speltoides induces homoeologous recombination, and thus will induce recombination between the chromosomes 4 Agropyron elongatum / 4T. monococcum or T. thaoudar / telomeric region of 4BL.
  • the final product must be a chromosome which combines the abovementioned genetic factors.
  • the phlb mutant or an equivalent method is used to induce homoeologues recombination.
  • a further technique designed to induce homoeologous recombination involves the use of the alien wheat species Triticum speltoides (Aegilops speltoides). When high pairing strains of T ⁇ speltoides are crossed (as the male parent) to wheat, the Phi gene is inhibited. This is the same affect as with the phlb mutant, ie, homoeologous recombination is expected to occur.
  • step 1 4thaoudar (or 4monoco ⁇ cum) is crossed with an addition line of Ag. elongatum, which has 43 chromosomes.
  • the progeny (F 1 ) is then crossed with the high pairing species T ⁇ speltoides, which is a monoploid species, and homoeologous pairing occurs in the offspring.
  • the critical F. is 20 + 4th +4E +4B +7S. It is expected that this genotype will occur at low frequencies.
  • step 3 the critical F 1 (20 + 4th + 4E + 4B) is crossed with +MS.
  • ms or ms.ms+i which is male fertile; note that ms.ms+i is a line homologous for male sterility carrying an isochromosome which confers male fertility. This line is used as a male donor of the ms gene because the isochromosome is transmitted through pollen at very low frequency.
  • step 4 the blue seed and fertile plants are selected to give the desired wheat.
  • EXAMPLE 7 The chromosome 4 substitution from Triticum monococcum also carries a height marker (gene for tallness). This is demonstrated in the results shown of the following table.
  • the short arm of 4m is known to carry the critical fertility restorer, if the height gene is present, also on this arm, male fertility will be initially identified by blue seed, and secondly by taller plants.
  • 91SU1 is a 4Ag blue addition line
  • 4th sub is a 4T. thaoudar substitution line
  • Embryos produced through the described procedures are routinely rescued by ⁇ ulturing the 15-20 day old embryos in special media, until a plantlet is produced. A brief summary of the procedure is described below.
  • the embryo culture methodology adopted involves the following steps: Step 1 : 15-20 days after crosspollination, excise seed from the inflorescence; Step 2 : wash seed 2 times in 6% sodium hypochlorite and sterile distilled water, and excise embryo from the endosperm; Step 3 : transfer embryo to a jar containing suitable media; Step 4 : wrap jar in foil, and leave at room temperature for 14 days; and Step 5 : transfer 2 leaf plantlet to plot. Plantlets derived from embryo rescue grow successfully.
  • the embryo rescue media used in this procedure is prepared as follows:

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Abstract

L'invention se rapporte à des plantes de blé à modification génétique et à la lignée obtenue de ces plantes, ainsi qu'à un procédé de production de blé hybride consistant à utiliser ces plantes. L'on obtient le blé génétiquement modifié en effectuant un appariement homologue pour ajouter un chromosome restaurateur de fertilité qui contient de la chromative télomère homologue à un autre chromosome de blé portant un gène récessif de stérilité mâle. Le chromosome restaurateur de fertilité contient une partie majeure de chromative étrangère provenant d'une autre espèce de blé, ainsi qu'un gène restaurateur de fertilité et au moins un gène marqueur. La structure génétique de la plante de blé, sous une forme préférée peut être décrite comme suit: 20'' + 4B(ms) + [4m (on 4th) - 4EL - 4BL]. Ces plantes présentent une stabilité méiotique et sont utilisées pour produire des souches de blé hybride.
PCT/AU1993/000017 1992-01-16 1993-01-18 Plantes de ble a modification genetique, lignee obtenue de ces plantes, et procede de production de ble hybride Ceased WO1993013649A1 (fr)

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AU33388/93A AU654762B2 (en) 1992-01-16 1993-01-18 Genetically modified wheat plants and progeny and method for production of hybrid wheat

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AUPL0436 1992-01-16
AUPL043692 1992-01-16

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WO1993013649A1 true WO1993013649A1 (fr) 1993-07-22

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Cited By (10)

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US5698686A (en) * 1994-10-20 1997-12-16 Arch Development Corporation Yeast telomerase compositions
WO1998051142A1 (fr) 1997-05-15 1998-11-19 Yeda Research And Development Co. Ltd. Procedes de production de ble hybride
WO1998056948A1 (fr) * 1997-06-10 1998-12-17 Pioneer Hi-Bred International, Inc. UTILISATION DE MARQUEURS MOLECULAIRES POUR LA DETERMINATION DU GENOTYPE DU GENE Rf OGURA CHEZ LES BRASSICA NAPUS
WO2002052924A3 (fr) * 2001-01-04 2003-02-20 Yeda Res & Dev Procede permettant de maintenir une ligne parentale genique male-femelle sterile pour la production de ble hybride
CN102835303A (zh) * 2012-09-14 2012-12-26 湖南杂交水稻研究中心 一种核质互作雄性不育系的育种方法
EP2918164A1 (fr) 2014-03-10 2015-09-16 Limagrain Europe Production de graines de céréales hybrides
US20160000030A1 (en) * 2013-03-18 2016-01-07 Sigrid WEISSMANN Methods of separating hybrid seed from a mixture of seeds
WO2019043082A1 (fr) 2017-08-29 2019-03-07 Kws Saat Se Aleurone bleue améliorée et autres systèmes de ségrégation
CN114457070A (zh) * 2022-01-30 2022-05-10 西北农林科技大学 一种小麦-二倍体长穗偃麦草45k液相芯片及应用
WO2023005883A1 (fr) 2021-07-28 2023-02-02 西南大学 Procédé de sélection de système de blé hybride à deux ligneés marqué en bleu et son utilisation

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AU6068890A (en) * 1989-08-10 1991-03-11 Bayer Bioscience N.V. Plants with modified flowers
AU8284891A (en) * 1990-07-17 1992-02-18 Pacific Seeds Pty. Ltd. Production of hybrid cereal crops

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Cited By (20)

* Cited by examiner, † Cited by third party
Publication number Priority date Publication date Assignee Title
US5698686A (en) * 1994-10-20 1997-12-16 Arch Development Corporation Yeast telomerase compositions
US5916752A (en) * 1994-10-20 1999-06-29 Arch Development Corporation Telomerase screening methods
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CN102835303A (zh) * 2012-09-14 2012-12-26 湖南杂交水稻研究中心 一种核质互作雄性不育系的育种方法
US10028460B2 (en) * 2013-03-18 2018-07-24 Bayer Cropscience Ag Methods of separating hybrid seed from a mixture of seeds
US20160000030A1 (en) * 2013-03-18 2016-01-07 Sigrid WEISSMANN Methods of separating hybrid seed from a mixture of seeds
US10743486B2 (en) 2013-03-18 2020-08-18 Basf Se Methods of separating hybrid seed from a mixture of seeds
EP2918164A1 (fr) 2014-03-10 2015-09-16 Limagrain Europe Production de graines de céréales hybrides
WO2019043082A1 (fr) 2017-08-29 2019-03-07 Kws Saat Se Aleurone bleue améliorée et autres systèmes de ségrégation
US20200255856A1 (en) * 2017-08-29 2020-08-13 KWS SAAT SE & Co. KGaA Improved blue aleurone and other segregation systems
US11697822B2 (en) 2017-08-29 2023-07-11 KWS SAAT SE & Co. KGaA Blue aleurone and other segregation systems
EP4591703A2 (fr) 2017-08-29 2025-07-30 KWS SAAT SE & Co. KGaA Ameliorations apportees a l'aleurone bleue et a d'autres systemes de segregation
EP4591704A2 (fr) 2017-08-29 2025-07-30 KWS SAAT SE & Co. KGaA Ameliorations apportees a l'aleurone bleue et a d'autres systemes de segregation
WO2023005883A1 (fr) 2021-07-28 2023-02-02 西南大学 Procédé de sélection de système de blé hybride à deux ligneés marqué en bleu et son utilisation
US12376536B2 (en) 2021-07-28 2025-08-05 Southwest University Breeding method and use of blue-grained two-line hybrid wheat system
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