WO2013190322A1 - Régulation de la dormance des graines - Google Patents

Régulation de la dormance des graines Download PDF

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Publication number
WO2013190322A1
WO2013190322A1 PCT/GB2013/051637 GB2013051637W WO2013190322A1 WO 2013190322 A1 WO2013190322 A1 WO 2013190322A1 GB 2013051637 W GB2013051637 W GB 2013051637W WO 2013190322 A1 WO2013190322 A1 WO 2013190322A1
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plant
nucleotide sequence
ubiquitin
ligase
nucleic acid
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PCT/GB2013/051637
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English (en)
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Steven Penfield
Sarah KENDALL
Dana MACGREGOR
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The University Of Exeter
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    • C—CHEMISTRY; METALLURGY
    • C12—BIOCHEMISTRY; BEER; SPIRITS; WINE; VINEGAR; MICROBIOLOGY; ENZYMOLOGY; MUTATION OR GENETIC ENGINEERING
    • C12N—MICROORGANISMS OR ENZYMES; COMPOSITIONS THEREOF; PROPAGATING, PRESERVING, OR MAINTAINING MICROORGANISMS; MUTATION OR GENETIC ENGINEERING; CULTURE MEDIA
    • C12N15/00—Mutation or genetic engineering; DNA or RNA concerning genetic engineering, vectors, e.g. plasmids, or their isolation, preparation or purification; Use of hosts therefor
    • C12N15/09—Recombinant DNA-technology
    • C12N15/63—Introduction of foreign genetic material using vectors; Vectors; Use of hosts therefor; Regulation of expression
    • C12N15/79—Vectors or expression systems specially adapted for eukaryotic hosts
    • C12N15/82—Vectors or expression systems specially adapted for eukaryotic hosts for plant cells, e.g. plant artificial chromosomes (PACs)
    • C12N15/8241—Phenotypically and genetically modified plants via recombinant DNA technology
    • C12N15/8261—Phenotypically and genetically modified plants via recombinant DNA technology with agronomic (input) traits, e.g. crop yield
    • C12N15/8262—Phenotypically and genetically modified plants via recombinant DNA technology with agronomic (input) traits, e.g. crop yield involving plant development
    • C12N15/8267—Seed dormancy, germination or sprouting
    • C—CHEMISTRY; METALLURGY
    • C07—ORGANIC CHEMISTRY
    • C07K—PEPTIDES
    • C07K14/00—Peptides having more than 20 amino acids; Gastrins; Somatostatins; Melanotropins; Derivatives thereof
    • C07K14/415—Peptides having more than 20 amino acids; Gastrins; Somatostatins; Melanotropins; Derivatives thereof from plants
    • C—CHEMISTRY; METALLURGY
    • C12—BIOCHEMISTRY; BEER; SPIRITS; WINE; VINEGAR; MICROBIOLOGY; ENZYMOLOGY; MUTATION OR GENETIC ENGINEERING
    • C12N—MICROORGANISMS OR ENZYMES; COMPOSITIONS THEREOF; PROPAGATING, PRESERVING, OR MAINTAINING MICROORGANISMS; MUTATION OR GENETIC ENGINEERING; CULTURE MEDIA
    • C12N9/00—Enzymes; Proenzymes; Compositions thereof; Processes for preparing, activating, inhibiting, separating or purifying enzymes
    • C12N9/93—Ligases (6)
    • C—CHEMISTRY; METALLURGY
    • C12—BIOCHEMISTRY; BEER; SPIRITS; WINE; VINEGAR; MICROBIOLOGY; ENZYMOLOGY; MUTATION OR GENETIC ENGINEERING
    • C12Y—ENZYMES
    • C12Y603/00—Ligases forming carbon-nitrogen bonds (6.3)
    • C12Y603/02—Acid—amino-acid ligases (peptide synthases)(6.3.2)
    • C12Y603/02019—Ubiquitin-protein ligase (6.3.2.19), i.e. ubiquitin-conjugating enzyme

Definitions

  • the disclosure relates to means for increasing seed vigour by providing a plant that expresses variants of ubiquitin E3 ligases and optionally variants of an ICE1 transcription factor in crop seeds and plants, and to crop plants and seeds comprising genetic variations of said gene(s) resulting in reduced/increased expression.
  • Methods enabling the identification of plants with modified expression of a ubiquitin E3 ligase and/or an ICE1 transcription factors.
  • a key feature of plant adaptive fitness is the ability to synchronise the onset of vegetative and reproductive development with seasonal changes in the environment.
  • the commencement of vegetative development is controlled by a period of quiescence in the mature seed known as seed dormancy.
  • seed dormancy a period of quiescence in the mature seed known as seed dormancy.
  • seed germination does not occur and the period of dormancy of many plant seeds is terminated by environmental signals including light, temperature and nutrient availability, a system adapted to the promotion of germination only when conditions are optimal for seedling establishment and reproductive success.
  • the role of light and temperature in the promotion of germination in dormant seeds is highly conserved among seed plants from angiosperms to gymnosperms, demonstrating the importance of germination control as a vital adaptive trait in plants.
  • Dormancy and seed germination is a hormonally regulated process and the major players are two phyto-hormones, abscisic (ABA) and giberrelic acid (GA).
  • ABA abscisic
  • GA giberrelic acid
  • ABA synthesis and GA catabolism is known to promote dormancy whilst GA synthesis and ABA catabolism stimulate dormancy breaking.
  • Genetic screens have also identified a number of loci such as ABI3, FUS3, LEC1 , DOG1 , RD04, MFT and FLC important in dormancy regulation.
  • CTR C-repeat
  • CBF1 -3 C-repeat-binding factor
  • ICE1 Inducer of CBF expression 1
  • HOS1 high expression of osmotically responsive gene 1
  • Loss of function hosl mutant plants showed enhanced cold induction of CBFs, and overexpression of CBFs confers freezing tolerance.
  • HOS1 overexpression confers increased sensitivity to freezing stress.
  • US2002/0148008 discloses the development of genetically modified wheat seed, in which the expression levels of Viviparousl (VP1 ) are modulated to regulate seed dormancy.
  • VP1 is a transcriptionally regulated gene essential for formation of seed dormancy; it is a transcription factor which acts in the ABA signalling system.
  • Site-directed mutagenesis of the gene results in a protein which comprises an amino acid sequence having deletions, substitutions or additions.
  • WO02/077163 describes the over expression of the gene ABI5 in plants such as Arabidopsis, to prevent precocious seed germination.
  • ABI5 encodes a putative transcription factor of the basic leucine zipper (bZIP) family.
  • bZIP basic leucine zipper
  • ABI5 has been shown to confer an enhanced response to exogenous ABA during germination.
  • ABA basic leucine zipper
  • As ABA-triggered processes, ABI5 protein accumulation, phosphorylation, stability and activity are highly regulated by ABA during germination and early seedling growth. Plants which over express ABI5 are hypersensitive to ABA and therefore respond to very low levels of this phytohormone, which would have no effect on wild type plants.
  • the disclosure relates to the identification of alleles of HOS1 and ICE1 for the identification of seed with high seed vigour, but minimising pre-harvest sprouting.
  • Methods for the production of such seeds and plants are also disclosed as the variation in the environment during seed production can have profound effects on the quality of seed produced for sale.
  • the mother plant has an important role in sensing the environment and using the information to modify the behaviour of seeds.
  • a genetic screen to identify Arabidopsis mutants with high seed vigour that is insensitive to the seed production environment.
  • HOS1 HIGH EXPRESSION OF OSMOTICALLY SENSTIVE GENES 1
  • a plant wherein said plant is not of the genus Arabidopsis and further wherein said plant expresses a nucleotide sequence variant of a gene comprising a nucleotide sequence that encodes a polypeptide that has ubiquitin E3 ligase activity and which plant produces seed that has increased vigour and/or low seed dormancy.
  • Seed vigour is defined as the ability of the seed to germinate at speed and establish with high frequency to form healthy strong-growing stands of plants under favourable and also less favourable environmental conditions. Seed as herein described with improved/increased seed vigour have at the time of germination a reduced response to sub-opitmal conditions such as temperature fluctuations, but still germinating efficiently and producing healthy seedlings. Low seed dormancy is an important determinant of vigour and low dormant seeds are less dependent of environmental factors which are, depending on the plant species, typically required to promote rapid germination.
  • sequence variant encodes an ubiquitin E3 ligase with reduced enzyme activity.
  • said plant is modified to provide an inactive ubiquitin E3 ligase.
  • said ubiquitin E3 ligase is encoded by a nucleotide sequence selected from the group consisting of: i) a nucleotide sequence as represented by the sequence in SEQ ID NO: 3, 4, 5, 6, 7, 8, 9, 1 1 ,12, 13, 14, 15, 16, 17, 18, 20, 21 , 22, 23, 24, 25, 26, 27, 28, 29, 30, 31 , 32, 33, 34, 35, 36, 37, 38, 39, 40, 41 , 42, 43, 44, 45 or 46;
  • nucleotide sequence wherein said sequence is degenerate as a result of the genetic code to the nucleotide sequence defined in (i);
  • nucleic acid molecule the complementary strand of which hybridizes under stringent hybridization conditions to the sequence in SEQ ID NO: 1 , 2, 10 or 19 wherein said nucleic acid molecule encodes a ubiquitin E3 ligase;
  • nucleotide sequence that encodes a polypeptide comprising an amino acid sequence as represented in SEQ ID NO: 49, 50, 51 , 52, 53, 55, 56, 57, 58, 59, 60, 61 , 62, 64, 65, 66, 67, 68, 69, 70, 71 , 72, 73, 74, 75, 76, 77, 78, 79, 80, 81 , 82, 83, 84, 85, 86, 87, 88, 89 or 90;
  • nucleotide sequence that encodes a polypeptide comprising an amino acid sequence as represented in SEQ ID NO: 47, 48, 54 or 63 wherein said amino acid sequence is modified by addition deletion or substitution of at least one amino acid residue as represented in iv) above and that has ubiquitin E3 ligase activity.
  • Hybridization of a nucleic acid molecule occurs when two complementary nucleic acid molecules undergo an amount of hydrogen bonding to each other.
  • the stringency of hybridization can vary according to the environmental conditions surrounding the nucleic acids, the nature of the hybridization method, and the composition and length of the nucleic acid molecules used.
  • the T m is the temperature at which 50% of a given strand of a nucleic acid molecule is hybridized to its complementary strand.
  • Hybridization 5x SSC at 65°C for 16 hours
  • Hybridization 5x-6x SSC at 65°C-70°C for 16-20 hours Wash twice: 2x SSC at RT for 5-20 minutes each
  • a modified polypeptide as herein disclosed may differ in amino acid sequence by one or more substitutions, additions, deletions, truncations that may be present in any combination with reference to SEQ ID NO: 47, 48, 54 or 63.
  • Preferred variants are those that vary from a reference polypeptide by conservative amino acid substitutions. Such substitutions are those that substitute a given amino acid by another amino acid of like characteristics.
  • the variant polypeptides have at least 30% identity, even more preferably at least 35% identity, still more preferably at least, 40%, 45%, 50%, 55%, 60%, 65%, 70%, 75%, 80%, 85%, 90%, 95% identity, and at least 99% identity with most or the full length amino acid sequence illustrated herein.
  • said ubiquitin E3 ligase is encoded by a nucleotide sequence as set forth in SEQ ID NO: 3, 4, 5, 6, 7, 8, 9, 1 1 ,12, 13, 14, 15, 16, 17, 18, 20, 21 , 22, 23, 24, 25, 26, 27, 28, 29, 30, 31 , 32, 33, 34, 35, 36, 37, 38, 39, 40, 41 , 42, 43, 44, 45 or 46.
  • said plant has enhanced expression of a variant gene comprising a nucleotide sequence that encodes an ICE1 transcription factor that regulates the expression of one or more transcription factors.
  • said ICE1 transcription factor comprises a nucleotide sequence selected from the group consisting of: i) a nucleotide sequence as represented by the sequence in SEQ ID NO: 91 , 92, 93, 94, 95, 96, 97, 98, 99, 100, 101 , 102, 103, 104, 105, 106, 107 or 108; ii) a nucleotide sequence wherein said sequence is degenerate as a result of the genetic code to the nucleotide sequence defined in (i);
  • nucleic acid molecule the complementary strand of which hybridizes under stringent hybridization conditions to the sequence in SEQ ID NO: 91 , 92, 101 , 104 or 105 wherein said nucleic acid molecule encodes a transcription factor that regulates the expression of one or more transcription factors;
  • nucleotide sequence that encodes a polypeptide comprising an amino acid sequence as represented in SEQ ID NO: 109, 1 10, 1 1 1 , 1 12, 1 13, 1 14, 1 15, 1 16, 1 17, 1 18, 1 19, 120, 121 , 122, 123, 124 or 125;
  • v) a nucleotide sequence that encodes a polypeptide comprising an amino acid sequence wherein said amino acid sequence is modified by addition deletion or substitution of at least one amino acid residue as represented in iv) above and which has transcription factor activity that regulates the expression of a transcription factor.
  • said transcription factor is encoded by a nucleotide sequence as set forth in SEQ ID NO: 91 , 92, 93, 94, 95, 96, 97, 98, 99, 100, 101 , 102, 103, 104, 105, 106, 107 or 108.
  • said plant is a transgenic plant engineered to reduce or abrogate the expression of said ubiquitin E3 ligase activity and optionally is engineered to enhance the expression of a transcription factor.
  • said transgenic plant is transformed with a transcription cassette wherein said cassette encodes all or part of a nucleotide sequence that encodes an ubiquitin E3 ligase and is adapted for expression by provision of at least one promoter operably linked to said nucleotide sequence such that both sense and antisense molecules are transcribed from said cassette.
  • said cassette is adapted such that both sense and antisense ribonucleic acid molecules are transcribed from said cassette wherein said sense and antisense nucleic acid molecules are adapted to anneal over at least part or all of their length to form a inhibitory RNA or short hairpin RNA.
  • a technique to specifically ablate gene function is through the introduction of double stranded RNA, also referred to as small inhibitory/interfering RNA (siRNA) or short hairpin RNA [shRNA], into a cell which results in the destruction of mRNA complementary to the sequence included in the siRNA/shRNA molecule.
  • the siRNA molecule comprises two complementary strands of RNA (a sense strand and an antisense strand) annealed to each other to form a double stranded RNA molecule.
  • the siRNA molecule is typically derived from exons of the gene which is to be ablated. The mechanism of RNA interference is being elucidated. Many organisms respond to the presence of double stranded RNA by activating a cascade that leads to the formation of siRNA. The presence of double stranded RNA activates a protein complex comprising RNase III which processes the double stranded RNA into smaller fragments (siRNAs, approximately 21 -29 nucleotides in length) which become part of a ribonucleoprotein complex.
  • the siRNA acts as a guide for the RNase complex to cleave mRNA complementary to the antisense strand of the siRNA thereby resulting in destruction of the mRNA.
  • said cassette is adapted to express an antisense RNA wherein said antisense RNA is adapted to anneal to a mRNA sequence that encodes a ubiquitin E3 ligase.
  • said inhibitory RNA or antisense RNA is designed with reference to a nucleotide sequence selected from the group consisting of: i) a nucleotide sequence as represented by the sequence in SEQ ID NO: 3, 4, 5, 6, 7, 8, 9, 1 1 ,12, 13, 14, 15, 16, 17, 18, 20, 21 , 22, 23, 24, 25, 26, 27, 28, 29, 30, 31 , 32, 33, 34, 35, 36, 37, 38, 39, 40, 41 , 42, 43, 44, 45 or 46 ;
  • nucleotide sequence wherein said sequence is degenerate as a result of the genetic code to the nucleotide sequence defined in (i); iii) a nucleic acid molecule the complementary strand of which hybridizes under stringent hybridization conditions to the sequence in SEQ ID NO: 1 , 2, 10 or 19 wherein said nucleic acid molecule encodes a ubiquitin E3 ligase.
  • said cassette is part of an expression vector adapted for expression in a plant cell.
  • said transcription factor is encoded by a nucleotide sequence which is part of an expression vector wherein said nucleotide sequence is operably linked to a nucleotide sequence comprising a transcription promoter.
  • said promoter confers constitutive expression on said transcription factor.
  • said promoter confers regulated expression on said transcription factor.
  • said regulated expression is tissue or developmental ⁇ regulated expression.
  • said regulated expression is inducible expression.
  • the nucleic acid molecule in the vector is under the control of, and operably linked to, an appropriate promoter or other regulatory elements for transcription in a plant cell.
  • the vector may be a bi-functional expression vector which functions in multiple hosts.
  • promoter is meant a nucleotide sequence upstream from the transcriptional initiation site and which contains all the regulatory regions required for transcription.
  • Suitable promoters include constitutive, tissue-specific, inducible, developmental or other promoters for expression in plant cells comprised in plants depending on design. Such promoters include viral, fungal, bacterial, animal and plant-derived promoters capable of functioning in plant cells.
  • Constitutive promoters include, for example CaMV 35S promoter (Odell et al. (1985) Nature 313, 9810-812); rice actin (McElroy et al. (1990) Plant Cell 2: 163-171 ); ubiquitin (Christian et al. (1989) Plant Mol. Biol. 18 (675-689); pEMU (Last et al. (1991 ) Theor Appl. Genet. 81 : 581 -588); MAS (Velten et al. (1984) EMBO J. 3. 2723-2730); ALS promoter (U.S. Application Seriel No. 08/409,297), and the like.
  • Other constitutive promoters include those in U.S. Patent Nos. 5,608,149; 5,608,144; 5,604,121 ; 5,569,597; 5,466,785; 5,399,680, 5,268,463; and 5,608,142, each of which is incorporated by reference.
  • Chemical-regulated promoters can be used to modulate the expression of a gene in a plant through the application of an exogenous chemical regulator.
  • the promoter may be a chemical-inducible promoter, where application of the chemical induced gene expression, or a chemical-repressible promoter, where application of the chemical represses gene expression.
  • Chemical-inducible promoters are known in the art and include, but are not limited to, the maize ln2-2 promoter, which is activated by benzenesulfonamide herbicide safeners, the maize GST promoter, which is activated by hydrophobic electrophilic compounds that are used as pre-emergent herbicides, and the tobacco PR-1 a promoter, which is activated by salicylic acid.
  • promoters of interest include steroid-responsive promoters (see, for example, the glucocorticoid-inducible promoter in Schena et al. (1991 ) Proc. Natl. Acad. Sci. USA 88: 10421 -10425 and McNellis et al. (1998) Plant J. 14(2): 247-257) and tetracycline-inducible and tetracycline-repressible promoters (see, for example, Gatz et al. (1991 ) Mol. Gen. Genet. 227: 229-237, and US Patent Nos. 5,814,618 and 5,789,156, herein incorporated by reference.
  • tissue-specific promoters can be utilised.
  • Tissue-specific promoters include those described by Yamamoto et al. (1997) Plant J. 12(2): 255-265; Kawamata et al. (1997) Plant Cell Physiol. 38(7): 792-803; Hansen et al. (1997) Mol. Gen. Genet. 254(3): 337-343; Russell et al. (1997) Transgenic Res. 6(2): 157- 168; Rinehart et al. (1996) Plant Physiol. 1 12(3): 1331 -1341 ; Van Camp et al. (1996) Plant Physiol. 1 12(2): 525-535; Canevascni et al. (1996) Plant Physiol.
  • operably linked means joined as part of the same nucleic acid molecule, suitably positioned and oriented for transcription to be initiated from the promoter.
  • DNA operably linked to a promoter is "under transcriptional initiation regulation" of the promoter.
  • the promoter is a tissue specific promoter, an inducible promoter or a developmental ⁇ regulated promoter.
  • nucleic acid constructs which operate as plant vectors. Specific procedures and vectors previously used with wide success in plants are described by Guerineau and Mullineaux (1993) (Plant transformation and expression vectors. In: Plant Molecular Biology Labfax (Croy RRD ed) Oxford, BIOS Scientific Publishers, pp 121 -148.
  • Suitable vectors may include plant viral-derived vectors (see e.g. EP194809).
  • selectable genetic markers may be included in the construct, such as those that confer selectable phenotypes such as resistance to herbicides (e.g. kanamycin, hygromycin, phosphinotricin, chlorsulfuron, methotrexate, gentamycin, spectinomycin, imidazolinones and glyphosate).
  • a plant according to the invention for use in the improvement of seed vigour and/or seed dormancy.
  • a method for the preparation of F1 hybrid seed and the formation of an F1 hybrid plant comprising the steps: i) providing a first male sterile female plant cultivar wherein said plant encodes a plant ubiquitin E3 ligase variant and crossing said plant with a second male plant cultivar; and
  • said ubiquitin E3 ligase variant is inactive or has reduced enzyme activity.
  • said male sterile female plant is genetically male sterile.
  • said male sterile female plant is treated with an agent that induces male sterility.
  • Improving crop yield by crossing two inbred plant lines is a common plant breeding technique, which can result in heterosis.
  • Male plant sterility, the inability to produce viable pollen and so inhibit self-crossing, is often necessary to produce F1 hybrids efficiently on an economical scale.
  • plant male sterility there are several methods to introduce plant male sterility such as chemical hybridising agents such as 2-chloroethylphosphonic acid, sodium 1 -(p- chlorophenyl)-1 ,2-dihydro-4,6-dimethyl-2-oxonicotinate,3-(p-chlorophenyl)-6-methoxy-s- triazine-2,4 (1 H,3H) dione-triethanolamine, 2,7-diamino-10-ethyl-6-phenylantridium bromide, or utilising plant lines comprising genes causing cytoplasmic male sterility, genetic male sterility or cytoplasmic-genetic male sterility.
  • chemical hybridising agents such as 2-chloroethylphosphonic acid, sodium 1 -(p- chlorophenyl)-1 ,2-dihydro-4,6-dimethyl-2-oxonicotinate,3-(p-chlorophenyl)-6-methoxy-s- tria
  • a gene comprising a nucleic acid molecule that encodes a plant ubiquitin E3 ligase as a means to identify a plant with altered expression of said ubiquitin E3 ligase wherein said plant has improved seed vigour and/or seed dormancy.
  • a method to produce a plant that has altered expression of an ubiquitin E3 ligase comprising the steps of: i) mutagenesis of wild-type seed from a plant that expresses a ubiquitin E3 ligase;
  • nucleic acid molecule is analysed by a method comprising the steps of: i) extracting nucleic acid from said mutated plants;
  • Mutagenesis as a means to induce phenotypic changes in organisms is well known in the art and includes but is not limited to the use of mutagenic agents such as chemical mutagens [e.g. base analogues, deaminating agents, DNA intercalating agents, alkylating agents, transposons, bromine, sodium azide] and physical mutagens [e.g. ionizing radiation, psoralen exposure combined with UV irradiation].
  • chemical mutagens e.g. base analogues, deaminating agents, DNA intercalating agents, alkylating agents, transposons, bromine, sodium azide
  • physical mutagens e.g. ionizing radiation, psoralen exposure combined with UV irradiation.
  • said plant encodes an ubiquitin E3 ligase variant that has reduced ubiquitin E3 ligase expression and/or activity.
  • said ubiquitin E3 ligase is encoded by a nucleotide sequence selected from the group: i) a nucleotide sequence as represented by the sequence in SEQ ID NO:
  • nucleic acid molecule the complementary strand of which hybridizes under stringent hybridization conditions to the sequence in SEQ ID NO: 1 , 2, 10 or 19wherein said nucleic acid molecule encodes a ubiquitin E3 ligase.
  • said plant is analysed to determine expression of an ICE1 transcription factor variant that regulates expression of one or more transcription factors.
  • said transcription factor is selected from the group consisting of: i) a nucleotide sequence as represented by the sequence in SEQ ID NO: 91 , 92, 93, 94, 95, 96, 97, 98, 99, 100, 101 , 102, 103, 104, 105, 106, 107 or 108; ii) a nucleotide sequence wherein said sequence is degenerate as a result of the genetic code to the nucleotide sequence defined in (i);
  • a nucleic acid molecule the complementary strand of which hybridizes under stringent hybridization conditions to the sequence in SEQ ID NO: 91 , 92, 101 , 104 or 105 wherein said nucleic acid molecule encodes a transcription factor that regulates the expression of one or more transcription factors.
  • a screening method that detects the expression of an ubiquitin E3 ligase and/or an ICE1 transcription factor in a plant comprising the steps: i) providing an isolated sample from a plant to be assessed for expression and extracting nucleic acid from said sample;
  • said ubiquitin E3 ligase is encoded by a nucleotide sequence as set forth in SEQ ID NO: 3, 4, 5, 6, 7, 8, 9, 1 1 ,12, 13, 14, 15, 16, 17, 18, 20, 21 , 22, 23, 24, 25, 26, 27, 28, 29, 30, 31 , 32, 33, 34, 35, 36, 37, 38, 39, 40, 41 , 42, 43, 44, 45 or 46.
  • said ICE1 transcription factor is encoded by a nucleotide sequence as set forth in SEQ ID NO: 91 , 92, 93, 94, 95, 96, 97, 98, 99, 100, 101 , 102, 103, 104, 105, 106, 107 or 108
  • said method is a real time PCR method for the detection and quantification of a nucleic acid encoding all or part of a nucleotide sequence that encodes said ubiquitin E3 ligase and/or an ICE1 transcription factor.
  • a screening method that detects the expression of an ubiquitin E3 ligase and/or an ICE1 transcription factor in a plant comprising the steps i) providing an isolated sample from a plant to be assessed for expression;
  • said ubiquitin E3 ligase polypeptide is represented by the amino acid sequence set forth in SEQ ID NO: 49, 50, 51 , 52, 53, 55, 56, 57, 58, 59, 60, 61 , 62, 64, 65, 66, 67, 68, 69, 70, 71 , 72, 73, 74, 75, 76, 77, 78, 79, 80, 81 , 82, 83, 84, 85, 86, 87, 88, 89 or 90.
  • said ICE1 transcription factor polypeptide is represented by the amino acid sequence set forth in SEQ ID NO: 109, 1 10, 1 1 1 , 1 12, 1 13, 1 14, 1 15, 1 16, 1 17, 1 18, 1 19, 120, 121 , 122, 123, 124 or 125.
  • said isolated plant samples comprise an array of samples isolated from a plurality of plants to be tested for expression of said ubiquitin E3 ligase and/or ICE1 transcription factor nucleic acid or polypeptide.
  • said method further includes DNA sequencing of genomic DNA isolated from a plant variant obtained by said method[s] to determine a variant sequence encoding or controlling expression of a ubiquitin E3 ligase variant.
  • said method further includes DNA sequencing of genomic DNA isolated from a plant variant obtained by the method[s] to determine the variant sequence encoding or controlling expression of a ICE1 transcription factor variant.
  • the sequence variants may be modifications to the coding sequences to create variant proteins with altered activity.
  • the sequence variants may be modifications to expression control sequences [i.e. promoter sequences] which alter the expression of the respective genes.
  • a plant obtained by the method according to the invention comprises a viral vector that includes all or part of a gene comprising a nucleotide sequence that encodes an ubiquitin E3 ligase.
  • said gene is encoded by a nucleic acid molecule comprising a nucleic acid sequence selected from the group consisting of: i) a nucleotide sequence as represented by the sequence in SEQ ID NO: 3, 4,
  • nucleotide sequence wherein said sequence is degenerate as a result of the genetic code to the nucleotide sequence defined in (i);
  • nucleic acid molecule the complementary strand of which hybridizes under stringent hybridization conditions to the sequence in SEQ ID NO: 1 , 2, 10 or 19wherein said nucleic acid molecule encodes a ubiquitin E3 ligase.
  • a viral vector comprising all or part of a nucleic acid molecule according to the invention.
  • a viral vector according to the invention in viral induced gene silencing of a plant ubiquitin E3 ligase.
  • Virus induced gene silencing is known in the art and exploits a RNA mediated antiviral defence mechanism. Plants that are infected with an unmodified virus induces a mechanism that specifically targets the viral genome.
  • viral vectors which are engineered to include nucleic acid molecules derived from host plant genes also induce specific inhibition of viral vector expression and additionally target host mRNA. This allows gene specific gene silencing without genetic modification of the plant genome and is essentially a non-transgenic modification.
  • the invention is typically applicable to crop plants and includes cereals, oils seed plants, fruits and ornamentals.
  • the invention is applicable to, for example, corn (Zea mays), canola (Brassica napus, Brassica rapa ssp.), alfalfa (Medicago sativa), rice (Oryza sativa), rye (Secale cerale), sorghum (Sorghum bicolor, Sorghum vulgare), sunflower (helianthus annuas), wheat (Tritium aestivum), soybean (Glycine max), tobacco (Nicotiana tabacum), potato (Solanum tuberosum), peanuts (Arachis hypogaea), cotton (Gossypium hirsutum), sweet potato (lopmoea batatus), cassava (Manihot esculenta), coffee (Cofea spp.), coconut (Cocos nucifera), pineapple (Anana comosus), citris tree (Citrus spp.) cocoa (Theobroma cacao), tea (Camellia
  • plants of the present invention are crop plants (for example, cereals and pulses, maize, wheat, potatoes, tapioca, rice, sorghum, millet, cassava, barley, pea, and other root, tuber or seed crops).
  • Important seed crops are oil-seed rape, sugar beet, maize, sunflower, soybean, and sorghum.
  • Horticultural plants to which the present invention may be applied may include lettuce, endive, and vegetable brassicas including cabbage, broccoli, and cauliflower, and carnations and geraniums.
  • the present invention may be applied in tobacco, cucurbits, carrot, strawberry, sunflower, tomato, pepper, chrysanthemum.
  • Grain plants that provide seeds of interest include oil-seed plants and leguminous plants.
  • Seeds of interest include grain seeds, such as corn, wheat, barley, rice, sorghum, rye, etc.
  • Oil-seed plants include cotton, soybean, safflower, sunflower, Brassica, maize, alfalfa, palm, coconut, etc.
  • Leguminous plants include beans and peas. Beans include guar, locust bean, fenugreek, soybean, garden beans, cowpea, mungbean, lima bean, fava bean, lentils, chickpea, etc.
  • Figure 1 shows that in Arabidopsis thaliana, hosl mutants have a strong germination vigour even when the temperature during seed production is lowered and the germination of wild type seeds is strongly inhibited. Therefore deleting HOS1 in plants causes germination vigour that is less sensitive to the temperature conditions during seed production;
  • FIG. 2 shows that HOS1 acts through the genome of the mother plant to control seed vigour
  • Figure 3 shows that hormone sensitivity in mature imbibed seeds is not very different in hosl mutants compared to wild type. This is consistent with HOS1 acting during seed production to control vigour;
  • Figure 4 shows that the seed coats of hosl mutants are more permeable to the metabolic dye tetrazolium. Permeability is known to be controlled by the biochemistry of tannins in the seed coat which is a tissue derived from the mother plant;
  • Figure 5 shows that the gene expression of enzymes in the tannin biosynthetic pathway is lower in hosl mutants than wild type, and that decreasing the temperature during seed production increases seed tannin biosynthetic gene expression. This is consistent with temperature and HOS1 acting to control seed coat biochemistry. Because seed coat tannin content is known to be associated with vigour, this represents a mechanism trherough which HOS1 may control seeed vigour from the maternal genome;
  • Figure 6 describes the germination vigour of mutants of the Arabidopsis thaliana ICE1 locus.
  • the HOS1 protein is known to interact physically with ICE1 in plant cells, and hosl mutants are known to have increased levels of the ICE1 protein.
  • ICE1 loss-of-function mutants have low germination vigour and icel gain-of-function mutants have high vigour.
  • This data is consistent with the control of seed vigour by HOS1 being mediated by ICE1 ; and
  • Figure 7 shows that icel loss-of-function mutants have visibly darkened seed coats, consistent with increased tannin levels.
  • Seed dormancy assays Seeds were produced from plants growing at ⁇ ⁇ 6°C in a Percival growth cabinet under long days in white light. Germination of mature seeds took place at 22 °C on water agar plates in white light for 7 days. Data shown for germination experiments are mean and standard errors of five biological replicate seed batches (from 5 individual plants).
  • Hormone sensitivity assays Seeds were sown on water agar plates with the indicated concentration of abscisic acid (ABA) or the gibberellin biosynthesis inhibitor paclobutrazol (PAC; Greyhound Chromatography, Liverpool, UK). Seeds were then cold stratified for 3 days before germinating at 22 °C for 7 days and scoring germination freqeuency.
  • ABA abscisic acid
  • PAC gibberellin biosynthesis inhibitor paclobutrazol
  • Tetrazolium assays The tetrazolium staining protocol was based on Debeaujon et al. (2000 dx.doi. org/10.1 104/pp.122.2.403). Freshly harvested seeds were incubated in water or an 1 % (w/v) aqueous solution of 2,3,5-Triphenyltetrazolium chloride (Sigma-Aldrich cat# T8877- 5G) in 96-well plates in the dark at 30 °C. At 24, 48 and 72 hours after the start of incubation, aliquots of seed were transferred to acetate paper, the liquid was removed from them and the seeds were scanned using a tabletop scanner at 12000 pixel resolution.
  • 2,3,5-Triphenyltetrazolium chloride Sigma-Aldrich cat# T8877- 5G
  • FIG 1 shows that in Arabidopsis thaliana, hosl mutants have a strong germination vigour even when the temperature during seed production is lowered and the germination of wild type seeds is strongly inhibited. Therefore deleting HOS1 in plants causes germination vigour that is less sensitive to the temperature conditions during seed production.
  • Figure 2 shows that HOS1 acts through the genome of the mother plant to control seed vigour.
  • Figure 3 shows that hormone sensitivity in mature imbibed seeds is not very different in hosl mutants compared to wild type. This is consistent with HOS1 acting during seed production to control vigour.
  • Figure 4 shows that the seed coats of hosl mutants are more permeable to the metabolic dye tetrazolium.
  • Permeability is known to be controlled by the biochemistry of tannins in the seed coat which is a tissue derived from the mother plant.
  • Figure 5 shows that the gene expression of enzymes in the tannin biosynthetic pathway is lower in hosl mutants than wild type, and that decreasing the temperature during seed production increases seed tannin biosynthetic gene expression. This is consistent with temperature and hosl acting to control seed coat biochemistry. Because seed coat tannin content is known to be associated with vigour, this represents a mechanism trherough which HOS1 may control seeed vigour fromt he maternal genome.
  • Figure 6 describes the germination vigour of mutants of the Arabidopsis thaliana ICE1 locus.
  • the HOS1 protein is known to interact physically with ICE1 in plant cells, and hosl mutants are known to have increased levels of the ICE1 protein.
  • ICE1 loss-of-function mutants have low germination vigour and icel gain-of-function mutants have high vigour. This data is consistent with the control of seed vigour by HOS1 being mediated by ICE1 .
  • Figure 7 shows that icel loss-of-function mutants have visibly darkened seed coats, consistent with increased tannin levels.
  • HOS1 and ICE1 both act to couple seed vigour to the environmental conditions during seed production, and that by mutating either gene we can uncouple the process and produce seeds of predictably high or low vigour regardless of the environment during seed production.
  • This invention the manipulation of seed vigour by ICE1 and HOS1 , can be used in commercial seed production to increase germination vigour and to cause predictable vigour when seeds are produced at different sites or at the same site but in a varying climate.

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Abstract

Cette invention concerne des moyens pour augmenter la vigueur des graines par utilisation d'une plante qui exprime des variants d'ubiquitine E3 ligases et éventuellement des variants d'un facteur de transcription ICE1.
PCT/GB2013/051637 2012-06-22 2013-06-21 Régulation de la dormance des graines WO2013190322A1 (fr)

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Cited By (5)

* Cited by examiner, † Cited by third party
Publication number Priority date Publication date Assignee Title
CN106244596A (zh) * 2016-08-29 2016-12-21 南京农业大学 一种水稻种子休眠基因OsDOG1L3及其应用
CN112760331A (zh) * 2020-11-13 2021-05-07 中国农业科学院作物科学研究所 大豆GmHOS1a基因和GmHOS1b基因及其应用
CN112778408A (zh) * 2021-03-02 2021-05-11 中国热带农业科学院橡胶研究所 橡胶树转录因子HbICE2及其编码基因与应用
CN116103309A (zh) * 2022-11-22 2023-05-12 陕西省杂交油菜研究中心 油菜BnHOS1基因及其提高植物抗寒性中的应用
CN117070536A (zh) * 2023-10-18 2023-11-17 河南大学三亚研究院 拟南芥hos1基因在调控叶片衰老中的应用

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Cited By (5)

* Cited by examiner, † Cited by third party
Publication number Priority date Publication date Assignee Title
CN106244596A (zh) * 2016-08-29 2016-12-21 南京农业大学 一种水稻种子休眠基因OsDOG1L3及其应用
CN112760331A (zh) * 2020-11-13 2021-05-07 中国农业科学院作物科学研究所 大豆GmHOS1a基因和GmHOS1b基因及其应用
CN112778408A (zh) * 2021-03-02 2021-05-11 中国热带农业科学院橡胶研究所 橡胶树转录因子HbICE2及其编码基因与应用
CN116103309A (zh) * 2022-11-22 2023-05-12 陕西省杂交油菜研究中心 油菜BnHOS1基因及其提高植物抗寒性中的应用
CN117070536A (zh) * 2023-10-18 2023-11-17 河南大学三亚研究院 拟南芥hos1基因在调控叶片衰老中的应用

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