CN100510083C - Omega 3 fatty acid desaturase gene of halophyte, expression vector thereof, vegetable cell and plant transformed therewith - Google Patents
Omega 3 fatty acid desaturase gene of halophyte, expression vector thereof, vegetable cell and plant transformed therewith Download PDFInfo
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- CN100510083C CN100510083C CNB2004100242986A CN200410024298A CN100510083C CN 100510083 C CN100510083 C CN 100510083C CN B2004100242986 A CNB2004100242986 A CN B2004100242986A CN 200410024298 A CN200410024298 A CN 200410024298A CN 100510083 C CN100510083 C CN 100510083C
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Abstract
The present invention provides an omega 3 fatty acid desaturase gene and an expression carrier thereof and plant cells and a plant formed by the transformation of the gene. The excessive expression ofThe invention provides a halophytes omega 3 fatty acid desaturase gene and its expression vector and the said gene transformed cell and plant. The said gene expressed excessively in plants can increas the gene in a plant body can increase the content of an alpha-linoleic acid in a leaf, the capability of cold resistance, salt resistance and eumycete resistance of plants can be improved, and the nue the Alpha-linolenic acid content of leaf, can increase the cold-resistant, salt-tolerant and antimycotic ability, and can promote nutritious value of vegetable. That antisense suppression of the saitritive value of vegetables, etc. can be improved; the antonymous inhibition of the expression of the gene in the plant body can reduce the content of the alpha-linoleic acid and improve the capabilitd gene expression in vivo plant can reduce Alpha-linolenic acid content, and can promote the plant high temperature resistance. The invention relates to chimeric gene construction with eukaryon expresy of high temperature resistance of the plants. The present invention relates to the construction of a chimeric gene with a eukaryotic expression promoter and relates to host cells for converting the sion promotor, and chimeric gene transformed host cell and recombination and transformation method. At same time, the invention provides a method for cultivating transgenic plant in which the gene is chimeric gene and a method for recombination and conversion. Simultaneously, the present invention also provides a method for cultivating excessive expression or antonymously restricting the transgeniexpressed excessively or antisense suppressed. c plant of the gene.
Description
Technical field
The invention belongs to the molecular biology of plants field, relevant with vegetable fatty acid metabolism.Further say the full length cDNA sequence and the amino acid sequence coded thereof that the present invention relates to Suaeda salsa (Suaeda salsa) plastid omega-3 fatty acid desaturase (ω 3 fatty acid desaturase), the overexpression and the Antisense Suppression expression vector that comprise this cDNA sequence are with the method for this sequence plant transformed and generation transgenic plant.
Background technology
(Polyunsaturated fatty acids is the important component of film fat PUFAs) to polyunsaturated fatty acid, and is the precursor of signaling molecules such as prostaglandin(PG), hemoglutinin.Fatty acid desaturase is to participate in PUFAs synthetic class of enzymes, their catalysis on the hydrocarbon chain specific position, inserts pair keys reaction (Los and Murata, Biochim Biophys Acta, 1998,1394:3-15).The omega-3 fatty acid desaturase is a kind of membrane bound enzyme, but catalysis from the linolic acid to the alpha-linolenic acid (α-linoleic acid, conversion ALA) and hexadecadienoic acid are to conversion (Kodama et al, the Plant Mol Biol of hiragonic acid, 1997,33 (3): 493-502).
Alpha-linolenic acid is ubiquity in plant, is important component (Horiguch et al, the PlantCell Physiol of plant cell membrane fat and storage fat, 1998,39:540-544), in chloroplast membranes, alpha-linolenic acid and hiragonic acid can account for 80% of film fat total amount.Alpha-linolenic acid also be cell signal molecule jasmonic acid precursor (Mique and Browes, Plant PhysiolBiochem, 1998,36:187-197), the defensive raction of involved in plant (Farmer et al, Planta, 1998,206:167-174).Simultaneously, alpha-linolenic acid be pollen development (McConn et al, Proc Natl Acad Sci USA, 1997,94:5473-5477) and cell maturation (Horiguch et al, Plant Cell Physiol, 1998,39:540-544) necessary.
In animal, alpha-linolenic acid and derivative thereof, the EPA (eicosapentaenoic acid) that is called as ＂ DHA (docosahexaenoic acid) ＂ and DHA (docosahexaenoic acid) are the important component of all cells film.DHA and EPA be the precursor of regulating molecules such as prostaglandin(PG), tranexamic acid (Dyera and Mullen, FEBS Letters, 200l, 494:44-47), and can reducing cholesterol, bring high blood pressure down, suppress thrombosis, promote that brain grows.Owing to self can not generate, alpha-linolenic acid must absorb from food, be a kind of essential fatty acid (Dyera and Mullen, FEBS Letters, 2001,494:44-47).
The omega-3 fatty acid desaturase mainly is present in algae and the plant, nematode (Caenorhabditis elegans) FAT-1 is unique representative (Spychalla et al of this enzyme in the animal, Proc Natl Acad Sci USA, 1997,94 (4): 1142-1147), and applied for that in calendar year 2001 (patent No. is 6,194 to United States Patent (USP), 167, Browes et al).The gene desB of coding omega-3 fatty acid desaturase has cloned and has obtained among the cyanobacteria Synechocystis.In the higher plant, the omega-3 fatty acid desaturase is arranged in chloroplast(id) and endoplasmic reticulum, in chloroplast(id), be electron donor with the ferredoxin, and in endoplasmic reticulum, electron donor is cytochrome b5 (Shanklin and Cahoon, Annu Rev Plant Physiol Plant Mol Biol, 1998,49:611-641641).The structure of omega-3 fatty acid desaturase comprises that 4 are striden the film district, there are three in conjunction with the Histidine bunch (HDCGH HXXXXXHRTHH HHXXXXHVIHH) of iron and three hydrophilic area α, β, ε (Knipple etal in the tenuigenin side, Genetics, 2002,162:1737-1752).
The omega-3 fatty acid desaturase is cloned from rape the earliest and is obtained (Arondel et al, Science, 1992,258:1353-1355), from plants such as rape, soybean, purple perilla, Arabidopis thaliana, mung bean, castor-oil plant, corn, wheat, parsley, tobacco, paddy rice, capsicum, Limnanthes douglasii, clone respectively again later on and obtain.The expression of omega-3 fatty acid desaturase is subjected to light (Nishiuchi et al, Plant Mol Biol, 1995,29 (3): 599-609 in the plant; Horiguch et al, Physiol Plant, 1996,96:275-283; Nishiuchi et al, Plant Cell, 1997,9 (10) 1701-1712) low temperature (erberich et al, Plant Mol Biol, 1998,36 (2): 97-306) injury (ibson et al, Plant Physiol, 1994,106:1615-1621; Kwon et al, Mol Cells, 2000,10 (5): 4997; Nishiuchi et al, Plant Cell, 1997,9 (10): 1701-1712), salt (Browse et al, J Biol Chem, 1993,268:16345-16351), growth hormone (Yamamoto, Planta, 1994,192:359-364; Matsuda et al, Planta, 2001,213:833-840), dormin (Zou et al, Plant Physiol, 1995,108:563-571), fungi infestation and peptide exciton Pep25, cycloheximide (Kirsch et al, Proc Natl Acad SciUSA, 1997,94 (5): the influence of the factor such as 2079-2084).
Plant omega-3 fatty acid delta 8 desaturase genes Arabidopis thaliana (Arondel et al, Science, 1992,258,1353-1355), tobacco (Hamada et al, Plant Physiol, 1998,118 (2): 591-598; Kodama et al, Plant Physiol, 1995,107 (4): 1177-1185), paddy rice (Kodama et al, Plant Mol Biol, 1997,33 (3): 493-502; Shimiada et al, PlantBiotechnol, 2000,17:43-48), sweet potato (Wakita et al, Plant Cell Reports, 200l, 20:244-249), Radix Dauci Sativae (Yadav et al, Plant Physiol, 1993,103 (2): 467-476) wait overexpression in the plant, and nematode fatty acid delta 8 desaturase genes overexpression in Arabidopis thaliana, all increase the content of alpha-linolenic acid, strengthened the plant resistance to cold.Transgene silencing (Maracari et al, Science, 2000,287:476-479) and Antisense Suppression (Hamada et al, Transgenic Res, 1996,5:115-121) the tobacco of plant omega-3 fatty acid delta 8 desaturase genes, the content of alpha-linolenic acid reduces, and can better adapt to high temperature.
Obtain the omega-3 fatty acid delta 8 desaturase genes though in various plants, cloned, but still do not find the patent of plant plastid omega-3 fatty acid delta 8 desaturase genes by the computer networking retrieval, associated patent is a United States Patent (USP) 6,194,167 (Browes et al, 2001), its omega-3 fatty acid delta 8 desaturase genes of cloning comes from the animal nematode, this assignment of genes gene mapping is on endoplasmic reticulum, its effect mainly is the synthetic of alpha-linolenic acid in the non-photosynthetic organ of catalysis (as seed, root etc.) in plant, and this is little with the relation that plant tolerates adverse circumstance.Still there is not at present report from the omega-3 fatty acid delta 8 desaturase genes of halophytes.
Plant will meet with various environment-stress inevitably in its life history,, illumination unbalance, too high or too low for temperature as mineral element excessively or deficiency, moisture shortage etc., these environment-stress influence the biochemical reactions of plant, cause plant-growth slow, having a strong impact on the output of agricultural and the quality of farm crop.According to estimates, the underproduction of farm crop is over half is in the world at present caused by environment-stress.
Summary of the invention
One of purpose of the present invention provides the gene that has the fatty acid desaturase effect on a kind of ω of being coded in 3 positions; Two of purpose provides the overexpression and the Antisense Suppression expression vector of this gene; Three of purpose provides with the vegetable cell of this gene transformation and plant.
Purpose of the present invention can realize by following technical measures
(1) the omega-3 fatty acid delta 8 desaturase genes of halophytes of the present invention comprises the nucleotide fragments of (a) encoding amino acid sequence comprising the base sequence shown in SEQ ID NO:1 or SEQ ID NO:2 basically; Or comprise the aminoacid sequence shown in SEQID NO:3 basically; Or with (a) complementary nucleotide fragments.
(2) nucleotide fragments in and the recombinant plasmid of corresponding carrier for expression of eukaryon formation or the recombinant plasmid that constitutes with corresponding antisense expression vector.
(3) agrobatcerium cell of the recombinant plasmid transformed in the usefulness (2).
(4) with the agrobatcerium cell plant transformed cell in (3).
(5) utilize the direct conversion method plant transformed of particle gun cell with the recombinant plasmid in (2).
(6) host cell in above-mentioned (2), (3), (4) or (5), the polypeptide of generation has the sequence shown in the SEQ ID NO:3, and can generate corresponding omega-3 fatty acid by catalysis ω 6 lipid acid.
(7) part of a kind of transgenic plant or plant, excessive generation alpha-linolenic acid of its part or all of cell or alpha-linolenic acid content reduce.
(8) a kind of method that makes plastid omega-3 fatty acid desaturase overexpression in host cell, comprising: recombinant plasmid utilizes the direct transformed plant cells of particle bombardment; Or utilize the agrobacterium mediation method transformed plant cells.
(9) method expressed in host cell of a kind of Antisense Suppression plastid omega-3 fatty acid desaturase comprises that recombinant plasmid utilizes the agrobacterium mediation method transformed plant cells.
The present invention embodies the vital role of halophytes Suaeda salsa plastid omega-3 fatty acid delta 8 desaturase genes in anti-contrary coercing, and it can keep the particularly stability of thylakoid membrane of plant membrane lipid effectively.The invention provides the anti-method of a kind of new cultivation: be about to first plastid omega-3 fatty acid delta 8 desaturase genes and import various glycophytes by various transgenic methods from halophytes against crop, vegetables, ornamental plant new variety or new lines, particularly crop and ornamental plant, gene of the present invention overexpression in plant materials can increase the content of alpha-linolenic acid in the leaf, cold-resistant, salt tolerant and the fungal resistance of plant can be improved, and the nutritive value of vegetables etc. can be improved.This gene of Antisense Suppression can reduce the content of alpha-linolenic acid in the intravital expression of plant, can improve the ability of plant withstand high temperatures.The invention provides the structure of the mosaic gene that has the eukaryotic expression promotor, and the host cell that transforms this mosaic gene and the method for recombinant conversion are provided; Simultaneously, also provide the method for cultivating the transfer-gen plant of overexpression or this gene of Antisense Suppression.
The basic DNA recombinant technology that this invention relates to and the concrete experimental technique of molecule clone technology can be referring to " molecular cloning " (Sambrook et al, 1989), the sequence of the nucleotide fragments of involved coding Suaeda salsa plastid omega-3 fatty acid desaturase and relevant polypeptide are listed in sequence table, wherein:
SEQ ID NO:1 is the full length nucleotide sequence of coding Suaeda salsa plastid omega-3 fatty acid desaturase cDNA;
SEQ ID NO:2 is one section nucleotide sequence of open reading frame among the coding Suaeda salsa plastid omega-3 fatty acid desaturase cDNA;
SEQ ID NO:3 is the aminoacid sequence of nucleotide sequence among the SEQ ID NO:2;
Embodiment
The structure in embodiment 1 Suaeda salsa cDNA library and cDNA clone's separation and order-checking
The present invention is to be the structure that material carries out the cDNA library with Suaeda salsa seedling over-ground part.Grow the Suaeda salsa seedling in 4 weeks with the NaCl solution-treated of 400mM after 48 hours, extract total RNA of its over-ground part tissue with RNAgent Kit (Promega), separate its mRNA with MESSAGEMAKER Kit (GIBCO BRL), the Oligo-dT primer that has the XhoI cloning site that provides with cDNA Synthesis Kit (Stratagene) synthesizes cDNA first chain, synthetic then second chain, 5 of cDNA ' end is after RNaseH digestion, scabble or mend flat with the pfu archaeal dna polymerase, be connected with the adapter that has the EcoRI cloning site, pack into then between the EcoRI and XhoI cloning site of phage-zap express vector, make up phage library.After the packaging extract packing of library by GigapackIII Gold (Stratagene), ehec infection (Escherichiacoli) strain X LOLR, λ-zap express vector obtains pBluescript KS plasmid after shearing, phage library is converted into plasmid library.The picking mono-clonal is put forward plasmid DNA at random from the substratum that contains tsiklomitsin (10mg/L) and kantlex (25mg/L), checks order with the T3 sequencing primer of standard, obtains the sequence of cDNA5 ' end, i.e. ESTs (Expressed SequenceTaqs).By the construction cDNA library, obtain 1500 surplus an ESTs.
The separation and the evaluation of embodiment 2 Suaeda salsa plastid omega-3 fatty acid desaturase full length cDNA clones
Adopt the BLAST homology analysis, isolate the cDNAs of coding Suaeda salsa plastid omega-3 fatty acid desaturase.BLAST (comprises all nonredundancy GenBank CDS, the sequence with Protein Data Bank of three-dimensional structure, SWISS-PROT protein sequence database, EMBL and DDBJ database) to carry out a kind of mode that homology is searched in BLAST ＂ nr ＂ database.BLASTX (Gish and States, Nature Genetics, 1993,3:266-272) be a kind of a kind of mode that the 6 kinds of translation sequences and the Protein Data Bank of nucleotide sequence are compared that NCBI (NationalCenter of Biology Information) provides.P value (probability) is searched the homology degree that the cDNA sequence is compared with a certain sequence in the database, and the P value is big more, and its two homology is just big more.Learn relatively that by BLASTX the homology of the plastid omega-3 fatty acid delta 8 desaturase genes of this sequence and tobacco is the highest, is 80%.From the BLAST interpretation of result, learn that this is a full-length cDNA.Order-checking obtains the full length cDNA sequence of 2191 base pairs, see sequence table SEQ ID NO:1, identified on this basis its open reading frame (Open Reading Frame, ORF), see SEQ ID NO:2, and inferred its aminoacid sequence (SEQ ID NO:3).Encode 423 amino acid of a segment length, molecular weight of this gene fragment are that 48.5KD, iso-electric point are 9.04 polypeptide.
This ATG code of sequence ATG is positioned at sequence ＂ AACAATGGC ＂, with L ü tcke (EMBO J, 1987,6:43-48) the plant initiation codon subsequence of Tui Duaning is in full accord, proves that this sequence is the cDNA homing sequence really.TargetP Serverv1.01 software (http://www.cbs.dtu.dk/services/TargetP) analysis infers that this proteic Subcellular Localization is a chloroplast(id).Have 3 Histidine bunch (HDCGH in the deduced amino acid, HGWRISHRTHH, HHDIGTHVIHH), fully with three Histidine bunch (HDCGH, HXXXXXHRTHH, HHXXXXHVIHH) (Lopez Alonso et al that combine iron of vegetable fatty acid desaturase tenuigenin side, Biochem Syst Ecol, 2003,31:1111-1124) coincide, prove this gene Suaeda salsa plastid omega-3-aliphatic acid desaturase of encoding really.
The structure of embodiment 3 plant overexpression carriers and the expression in plant thereof
Involved in the present invention to plant expression vector be pCAMBIAL3301, for making the foreign gene forward insert the pCAMBIAL3301 carrier, earlier gene fragment is inserted intermediate carrier pRT101 (Topferet al with SmaI and EcoRI cloning site, Nucleic Acids Res, 1987,15:5890), utilize the two external source segments of downcutting in PstI site on the pRT101 again, insert the corresponding site of pCAMBIAL3301 carrier, reaction is spent the night under being connected 16 ℃, and the connection product carries out enzyme with corresponding restriction enzyme and cuts checking.With pCAMBIAL3301 recombinant plasmid transformed agrobacterium strains GV3101, incubated overnight agrobatcerium cell in 28 ℃ of YEB substratum (containing the 60mg/ml Rifampin) is containing stroke plate on the LB flat board of 60mg/ml Rifampin, with the checking of alkaline lysis upgrading grain.
With Agrobacterium-mediated Transformation Arabidopis thaliana (Arabidopsis thaliana cv.Columbia), method for transformation be vacuum infiltration method (Clough et al, Plant J, 1998,16:735-743).Be the transgenic line that obtains isozygotying, more than the continuous selfing three generations of transformant.The transgenic arabidopsis plant of isozygotying is carried out the salt tolerance analysis, and the result shows: contrast increase about 10% of the content of alpha-linolenic acid in the transgenic line blade; Transgenic line can be grown under 200mM NaCl condition and be solid; Transgenic line at low temperatures upgrowth situation significantly better than contrast.
Available leaf disc transformation method (Horsch et al, Science, 1985,227:1229-1231) or protoplast transformation method (vanden Elzen et al, Plant Mol Biol, 1985,5:149-154) transform dicotyledons, make the omega-3 fatty acid desaturase change oil crops such as rape, soybean, vegetables such as cucumber, tomato over to, forest such as willow, apple and other dicotyledons are by screening the transfer-gen plant that Herbicid resistant obtains having certain anti-cold-peace salt tolerance.
Monocotyledons often adopts the particle gun direct injection to transform, and also can transform by agriculture bacillus mediated direct transformation system.Transform Agrobacterium LBA4404 with pCAMBIA 3301 carriers that have foreign gene, transform farm crop then, or utilize particle bombardment to transform some farm crop such as paddy rice, corn, wheat etc. and turfgrass class, can obtain cold-resistant transgenic plant.
The structure of embodiment 4 Antisense Suppression plant expression vectors and the expression in plant thereof
The involved plant expression vector of this patent is pCAMBIAL3301, for making this carrier of the reverse insertion of foreign gene, earlier gene fragment is inserted intermediate carrier pRT101 (Topfer et al with XhoI and BamHI cloning site, Nucleic Acids Res, 1987,15:5890) in, utilize the two external source segments of downcutting in PstI site on the pRT101 again, insert the corresponding site of pCAMBIAL3301 carrier, the connection product carries out enzyme with corresponding restriction enzyme and cuts checking.In pCAMBIAL3301 recombinant plasmid transformed agrobacterium strains GV3101, incubated overnight agrobatcerium cell in 28 ℃ of YEB substratum (containing the 60mg/ml Rifampin) is containing stroke plate on the LB flat board of 60mg/ml Rifampin, with the checking of alkaline lysis upgrading grain.
With Agrobacterium-mediated Transformation tomato, rape or soybean, method for transformation be leaf disc transformation method (Horsch et al, Science, 1985,227:1229-1231), screen with weedicide.Be the transgenic line that obtains isozygotying, more than the continuous selfing three generations of transformant, can obtain the oil variety that alpha-linolenic acid content reduces, and can obtain having the transfer-gen plant of certain thermo-tolerance.
Sequence table
The information of SEQ ID NO:1
(a) sequence signature:
*Length: 2191 base pairs
*Type: nucleic acid
*Chain: two strands
*Topological framework: linearity
(b) molecule type: cDNA
(c) initial source: Suaeda salsa
(d) sequence description: SEQ ID NO:1
The information of SEQ ID NO:2
(a) sequence signature:
*Length: 1272 base pairs
*Type: nucleic acid
*Chain: two strands
*Topological framework: linearity
(b) molecule type: cDNA
(c) initial source: Suaeda salsa
(d) sequence description: SEQ ID NO:2
The information of SEQ ID NO:3
(a) sequence signature:
*Length: 423
*Type: amino acid
*Chain: strand
*Topological framework: linearity
(b) molecule type: peptide
(c) initial source: Suaeda salsa
(d) sequence description: SEQ ID NO:3
Claims (5)
1, the omega-3 fatty acid delta 8 desaturase genes of a kind of halophytes, the nucleotide sequence that it is characterized in that described gene is shown in SEQID NO:1 or SEQ ID NO:2.
2, a kind of recombinant plasmid that is made of carrier for expression of eukaryon is characterized in that described plasmid contains the described gene of claim 1.
3, use the agrobatcerium cell of the described recombinant plasmid transformed of claim 2.
4, with the agrobatcerium cell plant transformed cell described in the claim 3.
5,, it is characterized in that described vegetable cell transforms with the direct conversion method of particle gun with the vegetable cell of the described recombinant plasmid transformed of claim 2.
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| CN102260708A (en) * | 2011-07-15 | 2011-11-30 | 山东省农业科学院高新技术研究中心 | Thionin gene Rs-AFP1 plant binary expression vector and construction method and use thereof |
| CN107326018B (en) * | 2017-08-15 | 2020-03-10 | 南京晓庄学院 | Method for cultivating plant low temperature resistant gene and transgenic low temperature resistant plant |
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